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GENERAL FUNCTION OF THE GALL BLADDER 
FROM THE EVOLUTIONARY STANDPOINT 



BY 



FRANK W. GORHAM 

RESEARCH ASSISTANT, DEPARTMENT OF PHYSIOLOGY 
NORTHWESTERN UNIVERSITY 



AND 



ANDREW CONWAY IVY 

PROFESSOR OF PHYSIOLOGY, NORTHWESTERN UNIVERSITY 




ZOOLOGICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 
VOLUME XXII, NUMBER S 

JUNE 21, 1938 
PUBLICATION 417 



GENERAL FUNCTION OF THE GALL BLADDER 
FROM THE EVOLUTIONARY STANDPOINT 



BY 



FRANK W. GORHAM 

RESEARCH ASSISTANT, DEPARTMENT OF PHYSIOLOGY 
NORTHWESTERN UNIVERSITY 



AND 



ANDREW CONWAY IVY 

PROFESSOR OF PHYSIOLOGY, NORTHWESTERN UNIVERSITY 




A NATURAL - v ^ > 
HISTORY ^ 




ZOOLOGICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XXII, NUMBER 3 

JUNE 21, 1938 



PUBLICATION 417 



PRINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 






GENERAL FUNCTION OF THE GALL BLADDER 
FROM THE EVOLUTIONARY STANDPOINT 



BY FRANK W. GORHAM AND ANDREW CONWAY IVY 



It has long been known to anatomists that certain animals do 
not possess a gall bladder. The evidence offered has usually been 
from individual dissections, and the explanations of the absence 
have been rather unsatisfactory. Therefore, an effort has been 
made to accumulate from the literature and dissections 1 a list of 
species in which the state of the organ is known. From this list 
certain physiological conclusions can be drawn. 

This work was stimulated by the physiological investigations 
of Schmidt and Ivy (1937) who studied domestic and wild animals, 
some with, others without, a gall bladder. They found that animals 
varied in the amount of bile produced by the liver, the size and con- 
centrating ability of the gall bladder, the motility of the common 
duct, and the resistance offered by the choledochoduodenal junction 
to the flow of bile into the intestine. From their data, they concluded 
that animals without a gall bladder produce large quantities of bile, 
while those with a gall bladder which concentrates well, produce 
small quantities of bile. Between these extremes lies a series of 
intermediate types, and, in general, it may be said that the amount 
of bile formed by the liver varies inversely with the efficiency of 
the concentrating apparatus associated with the biliary ducts. 
These observations suggested that by ascertaining the presence or 
absence of the gall bladder throughout the various vertebrate orders, 
evolutionary evidence might be obtained pertaining to the function 
of the organ. Also, information would become available, serving 
to further physiological and possibly genetic research. 

PHYSIOLOGICAL EVIDENCE 

To indicate the academic and practical significance of this 
problem, an outline of the physiology of the organ will be given. 

General Functions Assigned to the Gall Bladder. The general 
functions theoretically assigned to the gall bladder are those of serv- 
ing (a) as a bile reservoir for digestive purposes, and (6) as a pressure 

1 Field Museum of Natural History has been most generous in allowing the 
authors to use its collections and libraries. Without the aid of its staff members, 
this work could scarcely have been begun. We desire to express our thanks 
especially to Dr. Wilfred H. Osgood, Mr. Colin C. Sanborn, Mr. Karl P. Schmidt, 
and Mr. Rudyerd Boulton. 

159 



160 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

regulatory mechanism. These functions are suggested by the follow- 
ing evidence: (1) A sphincter is present at the junction of the common 
bile duct with the duodenum, and this sphincter is necessary for 
filling the gall bladder. When the sphincter is relaxed the gall 
bladder does not fill. (2) In some animals the secretory pressure 
of bile is markedly less than the pressure necessary to overcome the 
sphincter. To prevent injurious back pressure on the liver when 
the sphincter is forcefully contracted, a regulatory device is essential. 

(3) Cholecystectomy in animals that have a relatively powerful 
sphincter, leads in most instances to dilation of the common ducts. 

(4) Those animals which do not possess a gall bladder, physiologically 
have no sphincter or only a very weak one. Thus, when the gall 
bladder is present, a sphincter (Sphincter of Oddi; Sphincter ductus 
choledocus) is essential for its filling; and when an efficient sphincter 
is present a gall bladder is necessary for the regulation of biliary 
pressure. It is evident, however, that these observed facts bear 
more directly on pressure regulatory function than on that of bile 
storage for digestive purposes. 

Concentrating Activity of the Gall Bladder. In some animals, as 
man, the dog, and the cat, the storage function of the gall bladder 
is augmented by its ability to concentrate hepatic bile from five to 
ten times. In such animals, the gall bladder is literally a reservoir 
itself, of small volume but of large concentrating capacity. In other 
animals, as cattle, the gall bladder concentrates bile only twice or 
less. This ability of the organ to concentrate may be interpreted 
as supporting either or both of the aforementioned functions. 

Volume Output of Bile by the Liver. Animals vary widely in 
regard to the amount of bile secreted per kilo of body weight or 
per gram of liver. For example, the guinea pig secretes from 154 
to 220 cc. of bile per kilo of body weight per day. Man, however, 
only secretes 20 cc. of bile per kilo per day. 

Physiologic Capacity of the Gall Bladder. Since the size of the 
gall bladder, the bile output, and the ability of the gall bladder to 
concentrate varies in different animals, the physiologic capacity 
will vary. By this term is meant that fraction of the daily output 
of bile which the gall bladder is capable of storing. If it were known 
with certainty that this bile was stored for digestive purposes, this 
should be a part of the definition. A contrast in physiologic capacity 
is demonstrated by man, who can store the bile secreted during 12 
hours, and a guinea pig, which can store only the bile secreted for 
12 minutes. 



1938 GALL BLADDER GORHAM AND IVY 161 

An Additional Function of the Sphincter. It should be considered 
that this structure probably prevents the regurgitation of duodenal 
contents into the common bile duct. This prevents ascending in- 
fection of the liver. It suggests that animals devoid of a competent 
sphincter should have some compensatory mechanism. Several such 
mechanisms do occur. There may be a one-way valve; large quanti- 
ties of bile may keep the ducts flushed out; or the extra-hepatic 
ducts may show peristaltic activity. Investigations show that the 
horse, which lacks both a competent sphincter and a gall bladder, 
has both a valve and a liver, which forms large quantities of bile. 
On the other hand, the adult pigeon, 1 which also lacks a gall bladder, 
has motile ducts, and a liver forming large quantities of bile. 

Practical Significance of the Problem. It is not generally agreed 
that the gall bladder serves for the storage of bile for digestive 
purposes. This is because physiological studies have not as yet 
demonstrated detectable disturbances of digestion or nutrition after 
cholecystectomy. However, when a gall bladder which concentrates 
efficiently is removed, certain changes are known to occur. These 
are (a) incompetence of the Sphincter of Oddi, with a variable return 
of competence; (6) dilation of the extra-hepatic ducts; (c) small 
areas of local necrosis in the liver. These changes certainly indicate 
that the gall bladder has a function, even if its general role is not to 
store bile for digestive purposes. Unfortunately, it is not known 
why all these changes occur, but dilation of the ducts is generally 
attributed to the return of competence of the sphincter. If this 
interpretation is correct, and if no direct digestive disturbances 
result from cholecystectomy, then the chief function of the organ 
in man, the dog, and the cat, for example, is to regulate the pressure 
in the extra-hepatic ducts, and this regulation is rendered necessary 
by the Sphincter of Oddi. Hence, from an evolutionary viewpoint, 
a gall bladder was developed secondarily to the sphincter, and the 
sphincter served primarily to prevent regurgitation from the intestine. 
However, given an animal whose liver produces relatively small 
quantities of bile, it is then reasonable to assume that such an animal 
might possess a better digestive apparatus if a gall bladder and a 
sphincter were present to render possible the storage of bile for diges- 
tive purposes. The following facts force the consideration of such 
a hypothesis: (a) Bile plays an important role in the digestion and 
absorption of fats; (6) it is essential for the absorption of vitamin D 
and carotene, the precursor of vitamin A; (c) it is important in the 

1 A gall bladder is seen in the embryo. 



162 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

absorption of iron and calcium; (d) all domestic animals that form 
a small quantity of bile have a gall bladder of large physiologic 
capacity; (e) the absorption of bile salts from the intestine augments 
the secretion of bile by the liver. From this, it seems that a reserve 
supply of concentrated bile, which may be discharged into the in- 
testine in small amounts during the first 20 or 30 minutes after a 
meal, would have a favorable effect in initiating digestive processes 
in the intestine. In fact, the gall bladder bile is sometimes referred 
to by physiologists as "ignition bile." 

If it can be established that the storage of bile for digestive 
purposes is a function of the gall bladder, it will be obvious that 
cholecystectomized patients, or patients with chronic disturbances 
of the biliary tract, should be fed frequently with the idea of promot- 
ing the flow of bile by the liver, and thus keep it thin and moving 
as in animals where there is no gall bladder. That frequent feeding 
should be employed is an old clinical impression still held by some 
internists, but such management will not generally be recognized 
as essential until its physiologic importance is proved. For example, 
formerly it was debated whether fats and acid fruits should be 
avoided by the patient with acute cholecystitis. Now, since it is 
known that fats and acids are potent excitants of gall bladder activity, 
doubt no longer exists. 

In summary, the physiological evidence indicates that the chief 
function of the gall bladder is pressure regulation; but the possibility 
that storage of bile for digestive purposes also occurs in those forms 
which concentrate bile, is one which cannot be easily dismissed. 
While this remains unproved, it may be surmised that the gall bladder 
developed secondarily to the appearance of a sphincter at the junction 
of the common duct with the intestine. 

ANATOMICAL EVIDENCE 

Bearing the physiology of the gall bladder well in mind, it 
should be interesting to examine various vertebrate groups for the 
organ, to ascertain its presence or absence, and to correlate other 
facts pertaining to the economy of the animal. Thus, diet, other 
modifications of the digestive tract, and even taxonomic relationships, 
may bear on the existence of the organ in an individual species. 

Little effort has been made to describe variations in the form 
or anatomical relationships of the organ. Suffice it to say that this 
hardly seems necessary, since the organ is either present or absent, 
and no series between the extremes can be easily demonstrated. 



1938 GALL BLADDER GORHAM AND IVY 163 

"PREVERTEBRATES." Among lower forms, the liver has not 
consolidated and migrated to become an organ distinct from the gut. 
In Balanoglossus, the liver is represented by structures called "hepatic 
caeca," which are groups of cells surrounding numerous ductlike 
processes which connect with the alimentary tract. In the sea- 
squirts (ascidians), a group of glands communicates with the 
stomach by a duct. This is referred to as the digestive gland or 
liver. In Amphioxus the liver is similar. It is thus seen that none 
of the known predecessors of higher vertebrates possess an organ 
resembling a gall bladder. The gall bladder then becomes a typical 
vertebrate structure and very characteristic of the group. 

CYCLOSTOMATA. These are among the simplest vertebrates, but 
differ from all the others in lacking true jaws. The hagfish (Myxine) 
and the lamprey (Petromyzon) both have a gall bladder at some 
time during their lives. The adult hagfish has a bilobed liver, a 
gall bladder, but no discrete pancreas. The lamprey has a similar 
liver and gall bladder in the larval form, but the organ and its ducts 
are absent in the adult. 

FISHES. Recent fishes are divisible into six orders. Two of 
these, the sharks and rays (Elasmobranchii), and the chimaeras 
(Holocephali) have cartilaginous skeletons. The others have true 
bony skeletons. They are the sturgeon-like fishes (Chondrostei); 
forms related to the dogfish (Holostei); the true bony fishes 
(Teleostei); and the lungfishes (Dipnoi). The sharks and rays are 
representatives of a relatively primitive group which prospered 
during Devonian times. They all apparently possess a gall bladder. 1 
The lungfishes are interesting in that they represent a group similar 
to that from which Amphibia arose. They, too, retain the organ. 
The teleosts are the most successful of recent fishes and have been 
dominant since Cretaceous times. In them the gall bladder is 
occasionally absent and neither Cuvier (1835) nor Owen (1846) 
could offer any explanation for the variability. In those fish (Bull- 
heads) which have been studied, the gall bladder was found to con- 
tract and evacuate on the administration of fatty foods, but its 
concentrating ability is not known. 

AMPHIBIA. These all have a gall bladder in so far as they have 
been examined. This general statement is only superficially attested 
in the present work (Table 1). From the amphibians of the Coal 
Measures arose the reptiles which flourished during the Mesozoic, 

1 J. F. Daniel: The Elasmobranch Fishes. Univ. Calif. Press, 1922, p. 139. 



164 FIELD MUSEUM OF NATURAL HISTORYZOOLOGY, VOL. XXII 

and which gave origin during this period to the first mammals. 
Reptiles, therefore, are of current interest. 

REPTILES. All existing forms possess a gall bladder. Early 
workers reported its occasional absence, but their information was 
apparently erroneous. In Table 2 the presence of the organ is 
recorded in 42 families and 70 species. An attempt is made in this, 
as in subsequent lists, to select genera characteristic of their respec- 
tive families. 

It is perhaps unfortunate that no recent reptile approximates 
those from which mammals arose. But it is probable that these 
and, in fact, all reptiles possessed gall bladders. Therefore it is 
reasonable to believe that all early mammals likewise had the organ. 

The habit of intermittent feeding in reptiles is well known. 
Some of them are carnivorous and others herbivorous. These habits 
support the suggestion that the gall bladder is a reservoir of bile 
for digestive purposes. However, nothing is known concerning the 
physiology of the organ in this group. 

BIRDS. -This group appears to have been evolved from reptiles 
at about the same time as mammals, but it has attained a high 
degree of specialization and the number of the existing species is 
very large. 

With the available information, conclusions are difficult. 
Tendencies are perceptible, however. The carnivorous birds (hawks, 
owls, etc.) all retain the organ, while the herbivorous forms (parrots, 
pigeons) and the insectivorous forms (woodpeckers) may retain it 
or lose it. In no order yet examined is the organ invariably absent. 
It has been suggested (by R. Boulton, of Field Museum staff) that 
the insectivorous birds which have lost the organ were immediately 
derived from an old herbivorous ancestry. An example of interest 
here is the Kea parrot of New Zealand, whose carnivorous habits 
are as recent as the introduction of sheep into that region. 

The list of species in Table 3 is given as a matter of record, 
with the hope that its evident deficiencies will stimulate investigation 
by those who have more available material. 

MAMMALS. In this paper, mammals are of prime interest. This 
is largely because of man, and because it is more logical to cor- 
relate the physiology of man with that of other mammals than with 
that of some lower group. The orders will be considered separately. 

Monotremata. The monotremes are in many respects the most 
primitive of living mammals, yet in some respects they are highly 



1938 GALL BLADDER GORHAM AND IVY 165 

specialized. It seems not unlikely that they represent a line which 
descended directly, but independently, from the mammal-like 
reptiles. It may be recalled that they have a bill or beak devoid 
of teeth, and that they lay eggs but nourish their young on milk. 

The spiny anteater (Echidna) and the duck-bill (Ornithorhynchus) 
differ in their habits. One is terrestrial, and the other is semi- 
aquatic; one feeds on insects, and the other on shellfish and mollusks. 
Both have a gall bladder, and the cystic duct and the pancreatic 
duct join the common duct before it enters the intestine. Nothing 
is known concerning the physiology of the gall bladder or bile ducts. 

Marsupialia. Marsupials are considered higher than monotremes 
because they have abandoned the oviparous method of reproduction 
and now bring forth very immature young which are nursed in the 
marsupium, a structure characteristic of this group. The American 
opossum is a simple form which represents an ideal ancestor which 
has apparently passed almost unchanged from Cretaceous times. 
At the end of the Cretaceous, placentals began to appear and later 
became dominant. Somewhat previous to this, Australia with her 
mammalian fauna became isolated. With no other competition, 
marsupials there radiated adaptively until almost every ecologic niche 
was filled. Thus, we have marsupial forms which parallel super- 
ficially almost every other mammalian type. 

Food habits vary widely. The opossums (Didelphiidae) are 
essentially omnivorous. Forms like the Tasmanian wolf (Dasyuridae) 
are usually carnivorous, but other members of the family are in- 
sectivorous. The marsupial mole (Notoryctes) represents a mono- 
typic family (Notoryctidae) which is insectivorous. Species allied 
to the Koala (Phascolarctidae) are herbivorous and the honey- 
bear, itself, eats only the young shoots of a single species of Euca- 
lyptus. The wombats (Wombatidae) are herbivorous. The 
kangaroos and wallabies (Macropodidae) are herbivorous, and the 
large kangaroo is said to practice a kind of rumination. The interest- 
ing American marsupial, Caenolestes (Caenolestidae), is insectivorous 
with an omnivorous tendency. The anteater, Myrmecobius, is the 
lone representative of its family (Myrmecobiidae). 

As might be expected, the stomach of marsupials shows adapta- 
tions related to the diet. The stomach is simple in the zoophagous, 
entomophagous, and carpophagous forms. In those types which 
feed on the bulkier parts of plants, the stomach becomes complex. 
The gall bladder is present in all marsupials (Table 5) and is of large 



166 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

size, with no tendency to reduction in volume. Nothing is known 
concerning its concentrating ability. It contracts weakly, but the 
ducts and extraduodenal ampulla of the common duct are motile. 

The fact that all marsupials possess a gall bladder, even those 
which are strictly herbivorous, is of great interest. Among placental 
mammals, where the herbivorous type of diet has been assumed, 
the organ is absent in certain species. Remembering that marsupials 
are rather low in the scale of mammalian evolution, it would seem 
that the gall bladder is a primitive structure not easily lost. 

Table 5 affirms the presence of the organ in 26 species, representing 
the 10 existing families. 

Most living mammals belong to a more advanced group known 
as the placentals, in which a true placenta is constantly found. 
This structure permits a longer pre-natal period, and the need for 
the marsupial pouch is consequently lost. Other marked differences 
occur. 

Primitive placentals are known to have existed at the end of 
the Age of Reptiles. These primitive placentals were insectiv- 
orous, and from them arose the ancestors of our existing higher 
mammalian fauna. Members referable to the same order (Insecti- 
vora) as some of the early forms, still exist. 

Insectivora. Table 6 records the existence of the gall bladder 
in 34 species, representing all families. The name of the order 
implies the nature of the diet, but a number of species eat fresh 
flesh when it is offered. 

Chiroptera. Rather closely related to the insectivores, but of 
ancient and unknown origin are the bats. Bats have been very 
successful, and many families and species exist. They fall into 
two suborders, the Megachiroptera being the more primitive. 
The majority of these are large, and subsist on fruit and pollen. The 
stomach is rather complex. Most of the Microchiroptera are small 
and more highly specialized. The construction of the stomach 
differs according to the diet, which includes all types. Some 17 
families and about 2,000 different species and subspecies of bats are 
said to exist. Table 8 records the gall bladder present in 13 families 
and 59 species. Therefore it is very probable that it is present 
in all bats. 

Dermoptera. This order is represented by a single living genus, 
Galeopithecus (or Galeopterus). This animal, known as the flying 
lemur, is interesting, as it represents a form possibly similar to the 



1938 



GALL BLADDER GORHAM AND IVY 



167 



progenitors of bats. It is a fair-sized, herbivorous, tree-living form with 
a moderately complex stomach. The gall bladder is present (Table 7). 
Primates. Paleontologically and structurally, the primates are 
closely related to the primitive arboreal insectivores, the chief 
advances having been made in the development of a large brain, 
and stereoscopic vision. Three suborders are distinguishable. These 
are represented by the lemurs (Lemuroidea), by Tarsius (Tarsoidea), 
and by the true monkeys and apes (Anthropoidea). The lemurs 



::::: ORGAN PRESENT 
ORGAN ABSENT 
?>? ORGAN VARIABLE 




FIG. 1. Probable history of the gall bladder in placental mammals (adapted 
from Romer). 

are the most primitive of living primates, and they possess many 
insectivore affinities. Between the lemurs and the true monkeys is 
Tarsius, possessing some of the features of each group. The anthro- 
poids include monkeys, apes, and man. They are separable into the 
New World forms (Platyrrhines) and the Old World forms (Catar- 
rhines). The latter include the family Hominidae, of which man is 
the only member. 

The diet of primates is quite varied. As a group they are 
omnivorous. The more strictly herbivorous forms subsist chiefly 
on the more concentrated portions of plants, such as the seeds or 



168 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

fruit. The stomach of the lemurs and of Tarsius is simple. In the 
New World monkeys the stomach is simple except in the spider 
monkeys (Ateles), and the howlers (Alouatta), where there is some 
tendency to sacculation. In the Old World monkeys, the stomach 
is simple in the species which possess cheek pouches. In the langurs 
(Presbytis) and the guerezas (Colobu-s), which have no cheek pouches, 
the stomach is complex. In these monkeys, there is accumulation 
and retention of vegetable food in the stomach, and bezoars are 
not rarely found. The stomach reaches its maximum complexity in 
this group, which includes the sacred ape of India, one of the langurs. 
The gall bladder is present in representatives of all families of 
primates. Table 9 includes 68 species. 

Carnivora. Fossil evidence indicates clearly that all terrestrial 
fissiped carnivores arose from onecreodont family (Miacidae) probably 
since Eocene times. Eocene and Paleocene carnivores were known 
as creodonts and are directly descended from primitive insectivores 
of the preceding era. The ancestry of the marine forms is not known 
but is probably very similar. From the direct ancestry, diet, and 
feeding habits, these forms would be expected to possess constantly 
a gall bladder and this is well shown in Table 10 where 65 species 
representing all families were found to have the organ. 

Edentates. Under the general term "edentates" convenience 
may allow an inclusive discussion of three unique mammalian 
orders, the Xenarthra, Pholidota, and Tubulidentata. Linnaeus, 
Cuvier, and other early investigators noted the queer dental structure 
and the lack of relationships with other orders, and therefore placed 
all these together in a distinct but somewhat heterogeneous order. 
Careful examination of anatomical and paleontological evidence failed 
to reveal any true relationships, and separate orders were indicated. 

The order Xenarthra includes the armadillos, anteaters, and 
sloths. Existing species are only a shadow of a previously large and 
varied fauna. Recent forms are, for the most part, highly specialized. 
The anteaters subsist mostly on termites; the armadillos are insectiv- 
orous but feed frequently on carrion; the sloths are herbivorous. 
As in other orders, the stomach is adapted to the diet. A gizzard- 
like pylorus occurs in the insectivorous forms, while the stomach 
tends to become complex in the sloths. The gall bladder is present 
in all forms except Bradypus, the three-toed sloth. It is significant 
that variation in this order first appears in the herbivorous types. 
However, the question immediately arises as to why the organ exists 
in Choloepus, the two-toed sloth. Has this species been long distinct 



1938 GALL BLADDER GORHAM AND IVY 169 

or has it only recently diverged? The existing limited fauna with 
its paucity of species speaks against recent divergence and so also 
do marked structural differences other than the gall bladder. There- 
fore, Miller's (1923) classification is followed in which Choloepus 
stands as the only genus of a family. 

The order Pholidota, containing three genera, is the scaly ant- 
eater of the Old World. The two genera examined, Manis and 
Smutsia, possess a gall bladder and a gizzard-like pyloric antrum. 

OrycteropiLS is the single living representative of the order Tubu- 
lidentata. Its relationships are also unknown, but the work of Jepson 
(1932) suggests that the order was distinct by the end of the 
Eocene. The pyloric antrum is moderately thickened, and the animal 
possesses a gall bladder. 

The condition of the gall bladder among the Edentates is shown 
in Table 11. 

Artiodactyla. Existing artiodactyls form three distinct groups: 
(a) Suina: pigs, peccaries, hippopotamuses. (6) Tylopoda: camels 
and llamas, (c) Pecora: chevrotains, deer, giraffes, prongbucks, 
and bovids. These suborders were all separate by the end of the 
Eocene, but more primitive forms are found earlier in this epoch 
which give evidence as to the ancestry of the whole group. Matthew 
(1909) and Gregory (1910) believe that the Artiodactyla have been 
derived from unknown creodonts allied to the Mesonychidae. These 
are carnivorous, so it is perhaps correct to say that a cow is more 
closely related to a lion than it is to a horse. 

Among the Suina, the true pigs (Suidae) constantly possess the 
gall bladder. The peccaries (Tayassuidae) have lost the organ. As 
to the hippopotamus, the gall bladder was absent in one of the four 
recorded dissections. More data would be desirable. It is interesting 
that the pig has a simple stomach, while that of the hippopotamus 
and the peccary is complex. It has even been suggested that the 
two latter ruminate. 

The Tylopoda, camels and llamas, have long been distinct from 
other artiodactyls. They have complex stomachs and ruminate. 
They lack the gall bladder. 

All Pecora probably arose from forms similar to existing Traguline 
deer, the chevrotains, which have a gall bladder. True deer 
(Cervidae) have probably all lost the organ, with the exception of 
the musk-deer (Moschus). This Asiatic form represents a species 
which is transitory between the true deer and their Traguline 



170 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

ancestors. The retention of the gall bladder adds to the evidence 
afforded by the lack of horns and the excessive development of the 
canines, which indicates the primitive nature of this deer. 

The American prongbuck, not a true antelope, has a long 
separate history, and is the single living representative of the family 
Antilocapridae. It has a complex stomach and a gall bladder. 

The giraffes (Giraffidae) offer the curious possibility that the 
organ may be anomalously present in a very small percentage of 
cases. It has been reported in two of twenty recorded dissections. 

The* gall bladder is probably present in all members of the family 
Bovidae, with the exception of those in the subfamily Cephalophinae, 
a group of small African antelopes known as duikers in which it 
is constantly absent. The absence of the organ has been made a 
characteristic of this group by Pocock (1918). Crisp's (1862) report 
of its absence in two species of the subfamily Bubalinae should be 
rechecked in view of other errors made by this author. The report 
of the organ in two closely related groups by Garrod (1877) also 
needs confirmation. Tetracerus, a form which has long been juggled 
between the Cephalophinae and the Tragelaphinae, should by gall 
bladder evidence be classed with the latter, as is done by Weber 
(1927). 

In such an early distinct order, which has shown such great 
radiation and which has claimed so many now extinct genera, 
diversity of structure in existing species might be expected. The 
rather uniform presence or absence of the gall bladder in the various 
families lends significance to the gall bladder as an important struc- 
tural characteristic. Its occasional appearance in the giraffe and the 
primitive musk-deer is of interest in connection with their supposed 
Traguline origin. The retention of the organ in the Antilocapridae 
and most of the Bovidae lends support to the theory of the rather 
recent radiation of these families from the Tragulidae. The absence 
of the organ from the Cephalophinae may indicate a divergence of 
this subfamily from the Bovidae. At least, the presence or absence 
of the gall bladder follows quite well the taxonomy of this rather 
heterogeneous group (Table 12). 

"Sub-ungulates." Three orders of mammals are generally referred 
to under this name. They are the Hyracoidea, Proboscidea, and the 
Sirenia. Although existing species show few affinities, the earlier 
representatives had many similarities which suggest a common origin 
(Romer, 1936). 



1938 GALL BLADDER GORH AM AND IVY 171 

The Hyracoidea, known as conies or dassies, have one principal 
living genus (Procavia), confined to the Old World, and including 
various species. Superficially, they resemble rabbits, but dentition 
and foot structure unmistakably ally them to hoofed animals. Their 
stomach is rather complex and Lydekker reports that they ruminate. 
The Biblical l description is fitting: "Cheweth the cud, but divideth 
not the hoof." Records of dissections of these animals are confusing 
either because the gall bladder was variable, or the interpretation 
of the structure found was not uniform. The first explanation is 
probably correct. 

The order Proboscidea includes the elephants. These are highly 
specialized forms with a rather unique biliary anatomy. The gall 
bladder is absent but the common duct is wide, long, and has a 
reticulated mucosa. A large duodenal ampulla, called the terminal 
bile pouch, is present. It is divided irregularly into sacks and the 
pancreatic duct enters it. Owen (1866) says it is contractile. The 
structure is reminiscent of that found in some whales, and of 
the extra-duodenal ampulla of the guinea pig and opossum. 

The order Sirenia includes three recent families, one of which 
has become extinct in historic times. This last, Steller's sea-cow, 
apparently is the only one which lacks a gall bladder. This is 
difficult to explain, but the stomach is very complex. 

In animals which have evolved in the direction of these forms, 
a tendency to disappearance of the gall cyst is perceptible. In the 
elephants, this tendency is completely manifest; in the sirenia and 
hyraces it is less so. It seems that the latter are about to lose the 
organ. There is a pouch in the elephant which seems analogous to 
structures seen in some whales, but the elephant is a herbivorous 
type and the need for this compensatory structure is not obvious. 

Perissodactyla. This order, once more abundant, is now repre- 
sented by horses, tapirs, and rhinoceroses. In many respects, these 
forms have become highly specialized. It is well to point out that 
the order is an ancient one, first appearing at the base of the 
Eocene. When curiosity is expressed concerning the reason 
why two animals whose habits are as similar as those of the 
horse and the cow differ in respect to the gall bladder, it should 
be pointed out that they are of diverse ancestry. Thus, the 
gall bladder is constantly absent in the Perissodactyla (Table 14), 

1 Leviticus 11:5. 



172 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

but only occasionally so among the Artiodactyla. The stomach 
in this order is simple, but the immense size of the caecum is 
almost characteristic. In the wall of the common duct are some 
microscopic sacculi which may have compensatory significance. 
These are found in the horse but not in the tapir. 

Cetacea. Whales are highly specialized marine mammals of 
very wide distribution. Unlike most other aquatic mammals, they 
are incapable of any locomotion on land. Structural vestiges, how- 
ever, indicate that they were not always so confined and paleontolo- 
gists agree that they were probably derived from terrestrial carnivores 
some time in the Paleocene. By the end of the Eocene, they consti- 
tuted a distinct mammalian order. Thus, as were the horses so were 
the whales, exposed to evolutionary tendencies over a long period. 

Table 15 shows that the gall bladder was absent in all except 
one of the species recorded, and examination of this exception 
(Williams, 1838) leaves the impression that a mistake might well 
have been made. 

The absence of the gall bladder in the Cetacea is indeed odd, 
for these forms are highly carnivorous. They have very complex 
stomachs like ruminants, but the proventriculus is aglandular. The 
whale-bone whales (Mysticeti) feed on small marine invertebrates 
like cuttle-fish and squid; the toothed whales (Odontoceti) feed on 
fish and at least one genus, Orca, the killer, feeds on seals. The 
nature of the whale's diet makes continuous feeding highly improb- 
able. But the complex stomach and the observation that whole food 
is regurgitated when the animals are harpooned, led Beddard (1900) 
to suggest that the animals feed hurriedly and store their food in 
an accessory stomach. The digestion of this food would then be 
protracted over several hours. 

In Tursiops (Hein, 1915), and probably a good many other 
Odontoceti (Weissberg, 1933), a dilated bile reservoir is found in 
the course of the common duct. Into this drains the pancreatic 
duct, and between the reservoir and the intestinal outlet the common 
duct contains a valve similar to the valvula spiralis of the human 
cystic duct. This structure is very interesting and may represent 
a true compensatory mechanism. Were these animals of herbivorous 
ancestry, such biliary anatomy might be explained by suggesting that 
the whale was an animal which lost the gall cyst, changed its habits, 
and, when the need for the organ reappeared, developed the structure 
which is described above. With a proven carnivorous ancestry, 



1938 GALL BLADDER GORH AM AND IVY 173 

however, explanation is difficult, but it may be associated with the 
complex stomach, or a depth-pressure modification. 

Rodentia. Rodents are known from a long paleontological record, 
and they were early divisible into two suborders. One, Simplici- 
dentata, has two upper incisors; the other, Duplicidentata, has four. 
The rabbits and their allies comprise the latter group. 

More superficial divisions are quite natural. The squirrel-like 
forms, Sciuromorpha, are the simplest and the most primitive 
genera. Radiation at an early era resulted in our present diverse 
sciuromorph fauna, including six rather distinct families. 

Mouselike rodents, Myomorpha, are a more recent offshoot from 
early rodent lineage. Structural modifications indicate a degree of 
specialization not seen in the squirrel-like forms. Three families 
are included among the existing genera. The dormice, Myoxidae, 
retain a simplicity of structure indicating that forms similar to them 
may have been ancestral to the more highly specialized Dipodidae 
and Muridae. The jerboas, Dipodidae, are saltatorial forms of un- 
known ancestry. However, the greatest number of living rodents 
are found in the true rats and mice, the Muridae. Excepting man, 
they are the most successful of recent mammals, and, like man, their 
terrestrial range is practically unlimited. Radiation was apparently 
from the Old World. 

The other division of simplicidentate rodents, the Hystrico- 
morpha, includes a variety of forms of which the porcupine and the 
guinea pig are representative. The hystricid rodents are largely 
confined to South America, on which continent a great multiplicity 
of species has developed. 

The rabbits and hares have been placed in a distinct order 
(Lagomorpha) by many authorities. The possession of a pair of extra 
incisors was a differential point in Oligocene times, but morphological 
similarities probably best find recognition if a single order is used. 

Rodents are essentially herbivorous, but a number of them will 
accept a more omnivorous diet. The families may be considered 
separately: 

Aplodontiidae: These live near mountain streams and in dense 
vegetation of the Pacific northwest. They feed on various green 
plants. 

Sciuridae: These feed chiefly on nuts, seeds, and grass. They 
probably occasionally eat birds' eggs and insects. 



174 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

Castoridae: Beavers are aquatically adapted forms which fell 
trees. The bark of the trees is utilized for food. 

Heteromyidae: Kangaroo rats and mice of the Americas which 
live abundantly in arid, subdesert regions where plants bloom freely 
for only a few days during the year. The result is a large number 
of seeds. These are gathered and stored by the animals for future 
consumption. Apparently they hibernate. 

Geomyidae: Pocket gophers are fossorial and eat roots, bulbs, 
grass, and seeds. They may feed more or less constantly. They 
do not hibernate, but tunnel extensively under the snow when 
necessary. 

Pedetidae: These are fossorial and saltatorial forms, the Cape 
Jumping Hares. Their food seems to be entirely of a vegetable 
nature. 

Myoxidae: Arboreal and nocturnal creatures of small size. They 
feed on nuts and seeds, and probably hibernate. 

Dipodidae: Feed upon buds, leaves, twigs, and many kinds of 
plants; on seeds, grain, wild berries, chestnuts, acorns, grass, and bark. 

Spalacidae: The Cape Mole-rats live in subterranean burrows, 
which they dig in search for bulbs and roots. 

Muridae: This family includes a variety of rats and mice, whose 
habits were probably originally herbivorous. They easily modify 
their needs to the available supply. Their diet has in many cases 
become that of man, whom they parasitize. 

Bathyergidae: Members of this group are fossorial and subsist 
on a vegetable diet. 

Hystricidae: Old World porcupines whose food is entirely 
vegetable and consists mainly of roots. 

Erethizontidae: Arboreal forms, which eat the bark of trees. 
They do not hibernate. They are restricted to the New World. 

Dasyproctidae: The agoutis are tropical rodents, whose food 
consists of foliage, roots of ferns, fallen fruit, and possibly nuts. 

Caviidae: The best-known member of this family is the common 
guinea pig. In their natural state, the cavies feed on roots, corn, 
and other vegetable substances. 

Chinchillidae: Grass and roots form the chief substance of their 
diet. Long arid seasons may deprive these animals of water for 
considerable periods, but they seem to survive on the dried grass. 



1938 GALL BLADDER GORHAM AND IVY 175 

Capromyidae: Arboreal forms which live in the dense forest. 
Feed on fruits, leaves, and bark, but may also eat the flesh of small 
animals, particularly that of a kind of lizard. 

Octodontidae: The octodonts are represented by both African 
and South American variations. Both of these are herbivorous. 

Thryonomyidae: The African cane-rat which digs for roots and 
ground nuts. Extensive runway systems are formed under the grass 
and reeds. 

Ctenodactylidae: An African diurnal form. It lives among rocks 
and is herbivorous. 

Leporidae: The rabbits are diurnal and strictly herbivorous. 

Ochotonidae: These interesting mammals live in alpine rock 
slides. They gather grass which they stack and allow to dry. This 
supply keeps them during the winter months. 

It has been impossible to investigate thoroughly all the available 
rodent material, but a few noteworthy points may be mentioned. 
The stomachs of rodents vary markedly in their form and even in 
their histologic structure. This variation has been the subject of 
at least one paper. 1 Arrangement of rodent families into two groups, 
one with simple stomachs, and the other with complex stomachs, 
failed to reveal any correlation with the presence or absence of a 
gall bladder. 

The gall bladder is present in some families, absent in others, 
and variable in still others (Table 16). Further investigations are 
desirable and will probably alter these lists: 

Present 

Castoridae Spalacidae Capromyidae 

Anomaluridae Bathyergidae Thryonomyidae 

Myoxidae Dasyproctidae Leporidae 

Dipodidae Cayiidae Ochotonidae 

Chinchillidae 

Absent 

Aplodontiidae Heterpmyidae Petromyidae 

Geomyidae Pedetidae Erethizontidae 

Inconstant 

Sciuridae Hystricidae 

Muridae Octodontidae 

Conclusions are not accurate with such meager evidence. The 
whole order seems to be in a rather plastic state. Evolutionary 
tendencies are perceptible. 

The gall bladder is absent in the Aplodontiidae. In the Sciuri- 
dae it is usually present. The Pedetidae are intermediate between 

1 K. Toepfer. Die Morphologic des Magens der Rodentia. Morph. Jahrb., 17. 



176 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

the Sciuromorpha and Myomorpha and, as they are in other respects 
highly specialized, the gall bladder absence is not surprising. How- 
ever, the apparently close relationship which these forms have to 
the Anomaluridae, which are cholecystous, and the variation of the 
organ in the existing Myoidea, suggest an origin of this latter group 
from separate forms, some in which the organ was present and others 
in which it was absent. 

The geomyids and heteromyids are closely related and have been 
distinct since Oligocene times. 

Among the Myomorpha, the Myoxidae, Dipodidae, and the 
Spalacidae show simpler dentition than the Muridae. These three 
possess a gall bladder. The Muridae claims more genera than any 
other mammalian family and probably more individuals. In this 
family, the gall bladder is constantly present in some genera and 
absent in others. Its presence or absence may even be found in 
different species of the same genus. 

Some tendencies among the subfamilies are distinguishable. The 
Cricetinae, Lophiomyinae, and Hydromyinae usually do not possess 
the organ, while the Gerbillinae apparently tend to retain it. Among 
the Murinae proper, available records (nine genera) indicate vari- 
ability in this subfamily. The present distribution and the large 
number of species and subspecies show that these rodents are not 
only very successful, but that they are at a morphologically plastic 
stage in their development. The habits of the group and the variable 
presence of the gall cyst hint at its eventual disappearance. 

Hystricomorph rodents generally possess a gall bladder, but in 
a few instances the organ is absent. The New World porcupines, 
the Erethizontidae, all lack the organ, but it was present in six 
out of eight dissections of the Old World forms. The Erethizontidae 
have been distinct since the Oligocene. 

DISCUSSION 

From the above evidence, several facts are forthcoming. In 
the first place, it will be seen that the gall bladder is present in all 
carnivorous forms (except whales), and that it may be lacking among 
omnivorous or herbivorous forms. A carnivorous animal of neces- 
sity consumes only occasional meals, and these have a high fat 
content. From a physiological point of view, storage of concentrated 
bile should be a great aid in the digestion of such meals, and it is 
significant that the organ is constantly present in animals having 
these habits. 



1938 GALL BLADDER GORHAM AND IVY 177 

The absence of the organ in species of other dietary habits, is 
indicated by the evidence of Schmidt and Ivy. They found species 
in which the organ had no apparent or demonstrable function. It 
may be said that the organ was "physiologically absent." Species 
possessing this type of gall bladder are usually members of an order 
in which there are other species in which the organ is anatomically 
absent. Further, the gall bladder is not absent from those species 
which have descended most directly, or with the least specialization 
in regard to body form, from the ancestor of mammals. 

Another factor which may influence the existence of the gall 
bladder in a given species is the proximity of this species to the 
ancestral form. It will be noted that those species, families, or orders 
which most closely resemble the ancestor of a given group tend 
constantly to retain the gall cyst, while those which vary most, 
tend to lose the organ. The fact that other parts of the digestive 
tract adapt themselves quickly to dietary habits, suggests that the 
gall cyst may be the least plastic of the digestive organs. It is true, 
however, that it has completely disappeared in some mammalian 
families, whereas the stomach, caecum, and colon have not, although 
they frequently have undergone marked variations in form. 

The stable nature of the gall bladder suggests that it may have 
taxonomic value. In only nine families was the organ found to be 
present in some members and absent in others. In at least six of 
these the variation occurred between species or individuals. In the 
remaining three, it was possible to arrange subfamilies, and here, 
for the most part, definite tendencies following the lines of arrange- 
ment were demonstrable. Individual variations in the organ are 
probably comparable to the anomalies occasionally seen in the human 
cadaver. The frequency with which these anomalies occur may 
be a measure of the declining "need" for the organ. Thus in the 
dissections of Procavia, three observers found a gall bladder; five 
did not. On the other hand, with the giraffe the records show 
only two gall bladders out of about twenty dissections. From these 
records, it might be concluded that these animals will soon lose the 
organ completely. 

It is interesting that closely related species, particularly among 
the Muridae, may differ in the possession of the organ. Besides 
this, the occurrence of anomalous absences in individuals of the 
same species suggests that it might be possible to study the mechan- 
ism of the inheritance of the organ. 



178 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

CONCLUSIONS 

CD The gall bladder is a typical vertebrate structure, found first 
in larval agnathostomes and generally throughout the rest of the 
group. 

(2) All reptiles and amphibians have a gall bladder. 

(3) In birds the presence of the gall cyst is extremely variable, 
but carnivorous birds generally retain it. 

(4) The gall bladder is a primitive mammalian structure. It is 
not easily lost. It is not as readily lost as the form of the stomach, 
caecum, and colon is changed; i.e. a complex stomach may appear in 
a line of descent without the gall bladder's being lost. It is usually 
retained in forms which most closely resemble the ancestral type; 
forms which vary most widely may lose it. 

(5) Carnivorous mammals (except whales) possess a gall bladder, 
while mammals with other dietary habits may lose the organ. This 
apparently is related to the intermittent feeding habits of carnivorous 
types. Herbivorous forms with continuous feeding habits are most 
likely to lack the organ. 

(6) From the evidence of the dietary habits of cholecystous and 
acholecystous forms, one may conclude that if the gall bladder is 
removed from a given individual of any species, digestive efficiency 
will be diminished only if (a) the organ has a large physiologic 
capacity, and (6) intermittent meals of a high fat content are given. 

(7) There is no anatomical evidence to indicate that the Sphincter 
ductus choledochus developed before a gall cyst. Yet the presence 
of such a sphincter renders a gall bladder essential as a pressure 
regulatory apparatus, unless the common bile duct manifests definite 
peristaltic activity, as, for example, in the guinea pig. 



1938 GALL BLADDER GORHAM AND IVY 179 

TABLES 

Unless otherwise indicated all numbers are catalogue numbers of specimens in 
Field Museum. The presence of the gall bladder is indicated by + its absence 
by . The name in parentheses is the one used by the original author. Refer- 
ences to Gorham, 1936, are to Field Museum specimens dissected but not 
preserved. 

TABLE 1. AMPHIBIA 
Apoda 
Caecilidae 

+ Typhlonectes compressicauda ] 

+ Typhlonectes natans [ Fuhrmann 1914. 

+ Typhlonectes dorsalis J 

Salientia 
Pipidae 

+Pipa pipa (P. americana) Beddard 1895a. 

Pelobatidae 

-+Megophrys monlana (Xenophrys monticola) } 

+Megophrys hasseltii (Leptobatrachium) > Beddard 1907a. 

+Megophrys feae (Megalophrys) J 

Rhinophrynidae 

-\-Rhinophrynus dorsalis Giinther 1858. 

Leptodactylidae 

+Pseudis paradoxa (Rana) Hunter 1861. 

Microhylidae 

+Breviceps verrucosus Beddard 1908. 

Ranidae 

+Rana pipiens Gorham 1936. 

Caudata 
Necturidae 

-\-Megalobatrachus japonicus Beddard 1903. 

+Cryptobranchus alleghaniensis (Menopoma) 

Sirenidae 

+ Siren lacertina 



Amphiumidae 

+Amphiuma means (A. didactylum) . . . . 

Salamandridae 

-\-Salamandra salamandra (S. maculata) 



Hunter 1861. 



TABLE 2. REPTILIA 
Sauria 

Gekkonidae 

+Gekko gecko 8914. 

Eublepharidae 

+Coleonyx mitratus 5051. 

Pygopodidae 

+Lialis jicari 13869. 

Agamidae 

+Agama agama 19815. 

+Calotes mystaceus 14492. 

-i-Chlamydosaurus kingi Beddard 1905. 

Iguanidae 

-\-Crotaphytus collaris 637. 

+Phrynosoma blainvilli 8056. 

Cordylidae 

+Cordylus giganteus 19258. 

+Platysaurus guttatus 17322. 

-\-Chamaesaura aenea. . . . 17465. 



180 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 2. REPTILIA (Continued) 
Sauria (continued) 
Anguidae 

+Gerrhonotus infernalis 11200. 

+Anguis fragilis 22895. 

+Ophisaur us apus 1 5680. 

Anniellidae 

+Anniella pulchra Coe & Kunkel 1906- 

Helodermatidae 

+Heloderma suspectum Shufeldt 1890. 

Varanidae 

+ Varanus niloticus Beddard 1907. 

Xantusiidae 

-\-Lepidophyma flavomaculata 21788. 

Teiidae 

+Ameiva ameiva 16585. 

Amphisbaenidae 

-\-Amphisbaena alba 17802. 

Lacertidae 

+Lacerta viridis 15749. 

Gerrhosauridae 

+Gerrhosaurus nigrolineatus 13506. 

Scincidae 

+Eumeces schneideri 19636. 

Feyliniidae 

-\-Typhlosaurus vermis 16031. 

Dibamidae 

-\-Dibamus novae-guineae 14251. 

Chamaeleontidae 

-i-Chamaeleo pumilis (Chamaeleon) 1 

+Chamaeleo par Mob (Chamaeleon) .. : Beddard 190Tb> 

+Chamaeleo dilepis (Chamaeleon) i 

* Chamaeko verrucosus (Chamaeleon) i 

+Chamaeleo verrucosus (Chamaeleon) 18277. 

-\-Rhampholeon spectrum 19855. 

Serpentia 
Typhlopidae 

+ Typhlops punctatus 21081. 

Leptotyphlopidae 

-\-Leptotyphlops albifrons 87. 

Boidae 

+Charina bottae Cope 1898. 

+Constrictor constrictor 1 1404. 

+Sanzinia madagascarensis (Corallus) Beddard 1906a. 

Pythonidae 

+Python bivittatus 8925. 

+Python sebae Beddard 1904. 

Aniliidae 

+Anilius scytale 16943. 

+Anilius scytale (Ilysia) Beddard 1906. 

+ Anilius scytale (Boa) Owen 1833a. 

Beddard could not find the organ. 



1938 GALL BLADDER GORHAM AND IVY 181 

TABLE 2. REPTILIA (Continued) 
Serpentia (continued) 
Uropeltidae 

+Rhinophis blythii \ p e ters 1861. 

+Rhinophis oxyrhynchus ) 

Xenopeltidae / ^523 

+Xenopeliis unicolor | Thompson 1913. 

Achrochordidae 

+Chersydrus granulatus } 

Colubridae I Cope 1898. 

+Elaphe quadrivitlata 

+Heterodon contortrix J 

+Coluber constrictor Gorham 

Dasypeltidae 

+Dasypellis macrops 

Homalopsidae 

+Enhydris enhydris 

Boigidae 

-\-Boiga dendrophila 

Disteiridae 

+Pelamis platurus 

Elapidae 

+Naja nigricottis 

+Naja hannah Beddard 

+M icrurus fulmus 
Amblycephalidae 

+ Amblycephcdus moellendorffii 

Viperidae 

+ Vipera berus -17bb. 

Crotalidae 

+Crotalus riridis (C. confluentus) ^ O P ^ y 8 - 

+Crotalus horridus Owen 1833a. 

Testudinata 
Dermochelidae 

+Dermochelys coriacea Burne lyuo. 

Chelhydridae 

-rChelydra serpentina Martin 1830b. 

Testudinidae 

+Emys orbicularis (E. lutraria) Hunter 1 bl. 

+ Testudo elephaniina (T. indica) Martin 1 30. 

+ Testudo elephantina (T. elephantopus) Hunter 1 61. 

+ Testudo graeca Martin 1830a. 

+Geoemyda trijuga (Emys) j Anderson 1879. 

-f Kachuga ahongoka J 

+Clemmys leprosa j Kollman 1912. 

-\-Testudo mauritanica J 

Cheloniidae 

+Chelonia mydas (Chelone) Hunter 1861. 

Crocodilia 
Crocodilidae 

+Crocodylus cataphractus (C. leptorhyncus) Martin 183ob. 

+Crocodylus niloticus ("Crocodile du Nile") Geoffrey 1803. 

+Crocodylus acutus Owen 1831d. 

Rhynchocephalia 
Sphenodontidae 

+Sphenodon punctatus (Hatteria) Giinther 1867. 



182 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 3. AVES 
Struthioniformes 

Struthionidae f Garrod & Darwin 1872, 

-Slruihio camelus \ Hunter 1861. 

Rheiformes I Rothschild 1900. 

- Rheidae Rothschild 1900 

Casuariiformes 
Casuariidae 

+Casuarius ] 

Dromiceiidae | Rothschild 1900 

+ Dromiceius J 

-\-Dromiceius novae-hollandiae Boulart 1900 

Apterygiformes 
Apterygidae 

+Apteryx aiistralis Owen 1838 

Sphenisciformes 
Spheniscidae 

-\-Aplenodyles patagonica Reid 1835 

Procellariiformes 

+Procellariidae . . Forbes 1882a 

Pelecaniformes 
Phaethontidae 

+Phaethon lepturus (P. flavirostris) Beddard 1897a 

Pelecanidae 

+Pelecanus rufescens Martin 1835a 

-\-Pelecanus occidentalis (P. fuscus) 

-\-Pelecanus onocrotalus 



Phalacrocoracidae 

+Phalacrocorax carbo J 

Anhingidae 

+Anhinga anhinga (Plotus) Garrod 1876b. 

Ciconiiformes 
Ardeidae 

+Ardea cinerea. 



+Ardea purpurea (Nycticorax). 



Hunter 1861. 



+Botaurus stellaris. 

+Nyclicorax nycticorax (N. europaeus) . 
Cochleariidae 

+Cochlearius cochlearius (Cancroma) Murie 1867a. 

Balaenicipitidae f Beddard 1888. 

+Balaeniceps rex . . -j Fox 1929. 

Scopidae 1 Mitche11 1913 - 

+Scopus umbretta Beddard 1884. 

Ciconiidae 

+Leptoptilos crumeniferus Mentzer 1929. 

Phoenicopteridae 

-\-Phoenicoplerus ruber Hunter 1861. 

Anseriformes 
Anhimidae 

+Anhima cornida (Palamedea corniculaia) Beddard 1894. 



1938 GALL BLADDER GORHAM AND IVY 183 

TABLE 3. AVES (Continued) 
Anseriformes (continued) 
Anatidae 

-\-Cygnus olor \ u.. nt . or i o fi1 

+Branla canadensis (Anser) / Hunter 1 

+Branta bernicla Fox 1923. 

+Branta leucopsis \ u, . , aft1 

+Cairina moschata (Anser) / Hunter 1 

+Eulabeia indica Fox 1927. 

+ Mergellus albellus (Mergus) Kuhl 1820. 

Falconiformes 
Accipitridae 

+Gyps fulvus (Vultur) , 
+Aquila chrysaetos. 



+Haliaeetus albicilla 

+Accipiter nisus (Astur) 

+Accipiter gentilis (Astur palumbarius) 
4-Aviceda leuphotes 



Hunter 1861. 



.97002. 



Falconidae 

Falco peregrinus Kuhl 1820. 

Galliformes 
Cracidae 

+Penelope purpurascens (P. cristata) Hunter 1861. 

Phasianidae 

+Arborophila brunneopeclus 97003. 

Numididae 

+Numida meleagris Hunter 1861. 

Opisthocomidae 

Opisthocomus hoazin (O. cristatus) Young 1888. 

Gruiformes 
Turnicidae 

-Turnix tanki : 97007. 

Gruidae 

+Anlhropoldes virgo (Grus) Hunter 1861. 

Aramidae 

+Aramus scolopaceus Garrod 1876a. 

Rallidae 

-\-Porphyrio albus Hunter 1861. 

+Notornis mantelli Benham 1899. 

Heliornithidae 

Podica senegalensis Beddard 1890a. 

Rhynochetidae 

+Rhynochetos jubatus Murie 1867a. 

Cariamidae 

+Cariama cristatus (Dicholophus) Martin 1836. 

Otidae 

+Otis tarda Fox 1929. 

Charadriiformes 
Jacanidae 

+ Jacana spinosa (Parra) Forbes 1881a. 

Charadriidae 

+ Vanellus vanellus (V. vulgaris) . 



Scolopacidae I K ,, 

-Erolia alpina (Tringa) f Kutl 

-\-Calidris canutus (Arenaria calidria) I 

4- Numenius arqualu Hunter 1861. 

-\-Limnodromus griseus } 

Phalaropodidae [ Gorham 1936. 

+Lobipes lobatus J 



184 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 3. AVES (Continued) 
Charadriiformes (continued) 
Laridae 

+ Larus marinus Hunter 1861. 

\-Larus argentatus Hill 1926. 

+ Lanis ridibundus Kuhl 1820. 

Alcidae 

+ ('ria aalge (iroile) Hunter 1861. 

+ Uria aalge Kuhl 1820. 

+SynthUborhamphus antiques 1 g^g^t 1887 

+ Brachyramphus marmoratus / 

Columbiformes 

Pteroclididae 

+ Pterocles orientalis (P. arenarius) | Garrod 18741 

+ Syrrhaple8 Garrod 1874. 

Columbidae 

+Ptilinopus insolitus (Aedirhinus) \ u -.....-]] -i ooo 

+Coryphoenas crassirostris (Turacoena) / ^ 

Columba .... 
-Turtur 

Macropygia . 

Ectopistes . . . 

Chamaepelia. 

Metriopelia . . 

Zenaida 

Caloenas ... 

Didnnculus. . 

Chalcopelia . 



Tympanistria 



Garrod 1874. 



Ocyphaps. 

Leucosarda . . 

Phaps 

Phlogoenas . . 

Starnocnas . . 

Geopelia .... 

Goiira 

+Carpophaga . 
+Lopholaemus. 
-\-Ptilonopits . . 

Treron 

Columba vitiensis halmaheira (lanthoenas leucolaema) . . . . \ r . 187r; 
-Alectroenas pulcherrima (Erythroenas) / u 

Psittaciformes 

f Cuvier 1835. 

- Psittacidae -I Beddard 1898. 

{ Fox 1923. 

-\-Kakaioe goffini (Cacatua) 1 

+Kakaloe moluccensis (Cacatua) I /- arrof i isyyv, 

-\-Kakatoe haematnropygia (Cacatua philippinarum) [ 

+Nymphicus hollandicus (Calopsitta novae-hollandiae) . . . . j 
Cuculi formes 
Musophagidae 

+Gallirex porphyreolophus (Corythaix) Owen 1834. 

+ Turacuf persa (Corythaix buffoni) Martin 1836a. 

Cuculidae 

+Carpococcyx radiatus Beddard 1901a. 

+ Scythrops notae-hollandiae Beddard 1898a. 

+Cucnlus canorus Hunter 1861. 

+Ccnlropn* sinensis 97014. 



1938 GALL BLADDER GORHAM AND IVY 185 

TABLE 3. AVES (Confirmed) 
Strigiformes 

Strigidae 

+ Scolopelia peli Murie 1871. 

+ Phodilus badius Beddard 1890. 

+Asio otus (Otus aurifa) Hunter 1861. 

Caprimulgiformes 

Steatornithidae 

+Steatornis Beddard 1886a. 

Podargidae 

+Batracho8tomus Blyth 1866. 

+ Batrachostomus . 
+Podargus 

Aegothelidae 



+ Aegotheles . 



Beddard 1886a. 



Caprimulgidae 

-\-Caprimulgus 

Chordeiles sp 

+Chordeiles minor minor Gorham 1936. 

+Nyctidromus Beddard 1886a. 

Micropodiformes 
Trochilidae 

Archilochus colubris Crisp 1862a. 

Campylopterus ensipennis 97208. 

Trogoniformes 
Trogonidae 

+Harpactes erythrocephalus 97017. 

Coracii formes 
Alcedinidae 

Halcyon smyrnensis 97019. 

Meropidae 

+Melittophagus erythrocephalus 97187. 

Leptosomatidae 

+Leplosomu S discolor. . . { ^andld^and Milne-Edwards 1875. 

Bucerotidae 

+Bucorvns abyssinicus Garrod 1876. 

+Dichoceros bicornis (Buceros cavatus) Owen 1833. 

Piciformes 
Capitonidae 

+ Trachylaemus goffinii 97232. 

-\-Megalaema trirens 

+Cyanops franklinii (Megalaema) 

-\-Xaniholaema rosea 

Ramphastidae 



+Aulacorhynchus prasinus (Aulacorhamphus) . 



+Ramphasto8 discolorus 

+Ramphastos ritellinus 

-\-RamphasiGs piscivorous (R. carinatus) . 

-\-Pteroglossus aracari (P. wiedi) 

+Selenidera maculirostris 

Picidae 

Picumnus squamulatus 97216. 



Forbes 1882. 



186 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 3. AVES (Continued} 
Passeriformes 
Formicariidae 

-\-Myrmotherula schisticolor ..................................... 97214. 

Tyraniidae 

+M ecocerculus leucophrys ...................................... 97220. 

Pittidae 

-Pitta oatesi 
Philepittidae 

-PhilepHta ............................................ Forbes 1880b. 

Hirundinidae 

-\-Psalidoprocne fuliginosa .................................. .... 97260. 

Campephagidae 

+Pericrocotu8 flammeus ....................................... 97029 . 

Dicruridae 

^Chaptea aenea .............................................. 97032. 

Oriolidae 

+Oriolus chinensis ............................................ 97033. 

Corvidae 

+Chripsirhina temia ........................................... 97042. 

+Corvus brachyrhynchos ................................. Gorham 1936. 

+Pyrrhocorax pyrrhocorax ............................... Hunter 1861. 

Paridae 

Aegithaliscus annamensis ..................................... 97142. 

Timeliidae 

+Mesia argentauris ........................................... 97109. 

Alcippe nipalensis ......................................... < 



Mixornis rubricapilla ...................................... < Q^QOI' 

+Garrulax leucolophus ......................................... 97048. 

-\-Garndax vassali ............................................. 97045. 

+Stachyris nigriceps. . . . < Q-AKO' 

y < uoo. 
rycnonotidae 

+Criniger gularis ............................................. 97129. 

-\-Oiocompsa flanventris ...................................... / 97118. 

m j-j 97144. 

Turdidae ; 97149 

+Luscinia calliope .......................................... i Q"! 59 

Sylviidae { 97153. 

+Orlhotomus sutorius .......................................... 97186. 

Muscicapidae 

Siphia parva ................................................ 97166. 

+M uscicapula rubeculoides ..................................... 97172. 

+Culicicapa ceylonensis ....................................... 97163. 

Motacillidae 

+Anthus hodgsoni ............................................ 97186. 

Bombycillidae 

- Bombycilla garrula ...................................... Kuhl 1820. 

Laniidae 

+Laniarius atroflarus ......................................... 97257. 

+Lanius excubitor ...................................... Hunter 1861. 

Nectariniidae 

Cinnyris reichenom ......................... . < Q-OOO' 

^ y i -o-. 

Cinnyris jugularis ........................................... 97189. 

Aethopyga saturata ........... . .97194. 



1938 GALL BLADDER GORHAM AND IVY 187 

TABLE 3. AVES (Continued) 
Passeriformes (continued) 
Dicaeidae 

+Dicaeum concolor ............................................ 97201. 

Zosteropidae 

+Zosterop8 palpebrosa .......................... 

-\-Zosterops virens ............................................. 97284. 

Ploceidae 

+Estrilda melpoda ............................................ 97254. 

+Munia striata ............................................... 97205. 

Icteridae 

+Zarhynchus wagleri ......................................... 97229. 

Thraupidae 

+Calospiza chrysophrys guttala ................................. 97217. 

+Thraupis cyanocephala subcinerea ............................ 97218. 

Fringillidae 

+Allapele$ semirufus .......................................... 9721 1. 

TABLES 4-16. MAMMALIA 

TABLE 4 
Monotremata 

Ornithorhynchidae f Owen 1838. 

+Ornithorhynchus anatinus ............................. \ Crisp 1862. 

[ Flower 1872. 
+Ornithorhynchus analinus ("Platypus") ............... Mackenzie 1918. 



Tachyglossidae f 

+ Tachyglossus aculeatus (Echidna hystrix) ............. i Flower 187 9 

[ Chapman 1887. 
+ Tachyglossus aculeatus (Echidna sp.) .................. Mackenzie 1918. 

TABLE 5 
Marsupialia 

Didelphiidae [ Mackenzie 1918. 

-4-Didelphis marsupialis .......................... \ Hunter 1861. 

1 Flower 1872. 

+Didelphis paraguayensis (D. azarae) ................... { taws 834^*' 

+Didelphis virginianus .................................. Gorham 1936. 

+Monodelphis brevicaudatus (Didelphis hunteri) ............. Hunter 1861. 

Dasyuridae 

+Dasyurus quoll (D. tiverrinus) ....................... Mackenzie 1918. 

,. , . /Alston 1880. 

+Antechinomys lamger .............................. | Beddard 1908b. 

+ Thylacinu9 sp ........................................... Crisp 1862. 

+Phascogale tapoatafa (P. penciUata) ...................... Hunter 1861. 

-\-Sarcophilus harrisi ................................. Mackenzie 1918. 

Notoryctidae 

+Notorycies lyphlops ................................... Carlsson 1904. 

Paramelidae 

+Chaeropu castanotis ................................... Parsons 1903. 

Phalangeridae 

+Phalanger maculatus (Cuscus) ........................... Forbes 1881. 

+ Tricho8urus mlpecula (Phalangista mtlpina) ...... | j^ckenzieVgiS. 

+Pseudochirus sp .................................... Mackenzie 1918. 

-\-Schoinobale8 volans (Petaurus taguanoides) ................ Hunter 1861 



188 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 5 Continued 
Marsupialia (continued) 
Phascolarctidae [ Forbes 1881. 

-f Phascolarctos cinereus \ Sonntag 1921. 

Wombatidae I Mackenzie 1918. 

+ Wombatus ursinus (Phascolomys tvombat) Forbes 1881. 

+ Wombatus ursinus ("Wombat") Cleland 1869. 

+ Wombatus ursinus (Phascolomys sp.) Mackenzie 1918. 

Macropodidae / jj unter ^ggj 

-\-Macropus cangaru (M. major) < /-i _ , uco 

^ v_ nsp ioO . 

+Macropus cangaru (M. giganteus) Flower 1872. 

+Macropus parryi \ Mac k en zie 1918 

+M acropus walabatus J 

i-Megaleia rufa (Macropus ruber) Crisp 1862. 

+Dendrolagus inustus | Owvn 1852. 

-\-Dorcopsis luctuosa (Halmaturus luctuosus) Grarrpd 1875. 

+Dorcopsis luctuosa (Halmaturus xanthopis) Crisp 1862. 

+Polorous tridactylus (Hypsiprymnus murinus) Hunter 1861. 

Caenolestidae 

+Caenolestes obscurus Osgood 1921. 

Myrmecobiidae 

+Myrmecobius fasciata . . Mackenzie 1936. 

TABLE 6 
Insectivora 
Tenrecidae 

+Oryzorictes hova 1 nnh< , on 1 009 

+Centetes caudatus / Dobson 1882 - 

Potomogalidae 

*+Polomogale velox Seabra 1901. 

Solenodontidae 

+Solenodon paradoxus Allen 1910. 

+Atopogale cubanus (Solenodon) 1 

Chrysochloridae 

+Chrysochloris asiatica (C. aurea) [ Dobson 1882. 

+Chrysospalax dobsoni (Chrysochloris villosa) j 

+Amblysomus hottentotus (A. rutilans) j 

+Amblysomus obtusirostris (Chrysochloris) Peters 1852. 

Erinaceidae i Dobson 1881. 

+Erinaceus europaeus { piQ^L^J i^o ' 

{ Hill 1926.'" 

+Echinosorex albus (Gymnura rafflesi) Allen 1910. 

-+Paraechinus deserti (Erinaceus algirus) ) 

+Paraechinus micropus (Erinaceus pictus) 

-i-Paraechinus amir (Erinaceus macracanthus) 



+Paraechinus blanfordi (Erinaceus jcrdoni) 



Dobson 1882. 



-+Paraechinus niger . 

+ Hemiechinus grayi (Erinaceus grayi) 

-rAtelerix albiventris (Erinaceus) 

Aethechiiius angolae (Erinaceus diadematus) j 

+ Ericulu$ Mfairi (Echinops) Martin 1838. 

Soricidae 

i Sore x pulgaris Arnback 1907. 

+Sorex araneus Hill 1926. 

+Crocidura hirla Peters 1852. 

+ Blarina brevicauda 16545. 

Called "Vesicula vermiculair." 



1938 GALL BLADDER GORHAM AND IVY 189 

TABLE 6 Continued 
Insectivora (continued) 

Talpidae 





-\-Desmana moschata (Myogale) ......................... \ nnhann 

-f Scapanus latimanus (Scapanus townsendi) .............. J L 

Tupaiidae 

+ Tupaia belangeri ...................................... Garrod 1879. 

+Ptilocercus lowii ........................................ Clark 1926. 

+Dendrogale frenata ........................................... 46628. 

Macroscelididae 

+Rhynchocyon cirnei ................................... ] 

-\-\asilio brachyrhynchus (Macroscelides fuscus) ............ [ Peters 1852. 

+Petrodromus tetradactylus .............................. J 

+ Macroscelides rozeti ................... Duvernoy and Lereboullet 1840. 

TABLE 7 
Dermoptera 

Galeopithecidae 



+Galeopterus temmincki (Galeopithecus volans) ............ Chapman 1902. 

TABLE 8 
Chiroptera 

Pteropidae 

+Pteropus rufus rufus (P. edivardsi) ....................... Flower 1872. 

+Pteropus giganteus giganteus (P. medius) ................ \ _ K . , cc , 

+Pleropus subniger (P. rubicoUis) ........................ / KoDm l( 81 * 

+Pteropus edwardsi (P. edwardsii) ......................... Peters 1852. 

Cynopteridae 

*+Cynopterus brachyotis (C. scherzeri) 
*-\-Penthetor jagori (Cynopterus) . 



*-\-Epomops franqueti (Epomorphus comptus) 



Robin 1881. 



*+Hypsignathus monstrosus. 

*+Nyctymene cephalotes (Eonycteris) 

*+Roussettu$ amplexicaudaius (Cynonycteris) Robin 1881. 

+Rou$settus collaris (Cynonycteris) \ PQ , _, , flr - 9 

+Epomophorus crypturus / * 

Rhinopomidae 

+Rhinopoma microphyllum. 
Emballonuridae 



* -\-Emballonur a nigrescens. 



Robin 1881. 



*-(-/?/! ynchiscus naso (Rhynchonycteris) ................... 

* -\-Balantiopteryx plicata (Saccopleryx) ................... 

*+ Taphozus melanopogon .............................. 

+Coleura afra (Emballonura) .............................. Peters 1852. 

Xoctilionidae 

+Noctilio leporinus ...................................... Robin 1881. 

+Noclilio albiventer ...................................... Cuvier 1835. 

Nycteridae 



x Robin 1881. 

+ A yctens reiout ....................................... / 

+Xycteris hispida (N. villosa) ............................. Peters 1852. 

* The abeence of the gall bladder was not noted. 



190 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 8 Continued 
Chiroptera (continued) 
Megadermidae 

*+Megaderma spasma 1 

Rhinolophidae 



' '+Rhinolophus ferrum-equinum. 



Robin 1881. 



+Rhinolophus hipposideros . 
*+Rhinolophus euryale 

+Rhinolophus lobatus Peters 1852. 

Hipposideridae 

+ Hipposideros diadema (Phyllorhina) \ P..L; , aai 

*+Hipp08ideros commersoni (Phyllorhina) / * 

-\-Hipposideros commersoni (Phyllorrhina vittata) Peters 1852. 

* -\-Hipposider os armiger (Phyllorhina) , 
Phyllostomidae 

*+Phyllostomus hastatus 



*+Macrotus waterhousii. 



Robin 1881. 



*+Glossophaga soricina. 

*+Carollia perspicillata (C. brevicauda). . . 

*+Artibeus jamaicensis (A. perspicillatus) . 

+ Vampyrus spectrum Cuvier 1835. 

Desmodontidae 

+ Desmodus rotundus (D. rufus) Flower 1872. 

Vestertiliolidae 

-\-Vespertilio murinus. 

+Myolis myotis (Vespertilio) Kuhl 1820. 

-\-Myotis mystocinus (Vespertilio) } 

*+Scotophilus temminckii > Robin 1881. 

*+Pipistrellus kuhlii (Vesperugo) J 

+Pipistrellus pipistrellus (Vespertilio) Cuvier 1835. 

( Flower 1872. 

+Nyctalus noctula (Vesperugo) \ Hill 1926. 

( Cuvier 1835. 

*-}-Lasiurus borealis (Atalapha noveboracensis) j 

*+Barbastella barbastella (Synotus) [ Robin 1881. 

*+Plecotus auritus J 

+Plecotus auritus 1 P ia q- 

+Eptesicus serotinus ("Vespertillion") J 

+Eptesicus serotinus (Vespertilio) Kuhl 1820. 

+Scotophilus nigrita (Nycticejus planirostris) Peters 1852. 

*-\-M iniopterus schreibersii ) 

*+Kerivoula hardunckii 

Molossidae 

*+Af olossus obscurus 

*+Chaerephon plicatus (Nyctinomus) 

+Cheiromeles torquatus 

*+Tadarida brasiliensis (Nyctinomus) 

*+M ormopterus acetabulosus (Nyctinomus) 

+Nyctinomous limbatus (Dysopes) Peters 1852. 

+Nyctinomus taeniotus (Dinops cestoni) Cuvier 1835. 

TABLE 9 
Primates 
Lemuridae 

-\-Lemur mayottensis . . Beddard 1884a. 



> Robin 1881. 



+Lemur macaco Martin 183h 

* The aboence of the gall bladder was not noted. 



1938 GALL BLADDER GORHAM AND IVY 191 

TABLE 9 Continued 
Primates (continued) 
Lemuridae (continued) 

+Lemurcatta .......................................... Hunter 1861. 

+Lemur fulvus .......................................... Fl^ er }g?- 

+Lemurfuhus (L. albifrons) ............................. Hunter 1861. 

. , / Flower 1872. 
+ Lemur vanegatus (L. vanus) ..... j Grandidier and Milne-Edwards 1875. 

+Microcebus murinus .................................... Flower 1872. 

+Microcebus murinus (M. smithi) ........ ... ...... . . . . . ... .Huge iww. 

+Propithecus diadema .............. Grandidier and Milne-Edwards 1875. 

+Hapalemur griseus .................................. Bedda rd 1884a. 

+Hapalemur simus .................................... Beddard 1901. 

Indrisidae 

+Indris indris (Avahi laniger) ---- Grandidier and Milne-Edwards 1875. 



Daubentoniidae 

-\-Daubentonia madagascarensis (Chiromys) ..... Mj var t and Murie 1865. 



+Daubentonia madagascarensis ("aye-aye") ................. Owen 1866. 

Lorisidae 

+Loris tardigradus (Nyclicebus) ......................... . - Ruge 1902. 

/ Hunter 1861. 
+Loris tardigradus ................................... | Flower 1872. 

+Loris tardigradus (Nycticebus) ................. Mivart and Murie 1865. 

+Loris tardigradus (Stenops gracilis) ........................ Kuhl 1820. 

-\-Nycticebus javanicus ................................. j Flower 1872. 

+Arctocebus calabarensis ............................... J 

+Perodicticus potto .................................... | Flower 1872. 

-\-Galago madagascarensis .................................. Kuhl 1820. 

+ Galago crassicaudatus .................................. Flower 1852. 

+Galago mohali .......................................... Smith 1849. 

v / Beddard 1901. 
+Myoxicebus griseus (Hapalemur) . j Grandi dier and Milne-Edwards 1875. 

Tarsiidae ( Ruge 1902 

+ Tarsias fuscus (T. spectrum) ........................ \ Sonntag 1924. 

[ Woollard 



Callitrichidae 

+0edipomidas oedipus (Midas) .......................... ('^ W ^ T 186l' 

+Callithrix jacchus (Hapale) ........................... Beattie 1927. 

( r lower loTZ. 

r .., f Sonntag 1924. 

Cebldae Crisp 1862. 

+Cebus capucinus .................................... \ R uge I902a. 

I Flower 1872. 
+Cebus apella .......................................... Hunter 1861. 

+Ateks geoffroyi ........................................ Flower 1872. 

+Ateles cucullatus ....................................... Murie li 65. 

+Ateles ater ............................................. Ru ? e 1902a - 

+Ateles belzebuth ......................................... Kuhl l %' 

+Pithecia pithecia ...................................... Sonntag 1924. 

+Pitkecia monacha ...................................... 1?^!! JOQA 

+Cacajao rubicundus (Brachyurus) ........................ Forbes 1880. 

+Alouatta senicula ...................................... Flower 1872. 

+Saimiri sciurea (Callithrix sciureus) ...................... Martin 1833. 



192 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 9 Continued 
Primates (continued) 

Cercopithecidae , ... 109Q 

-\-Cercopithecus albogularis . . . . { Owenl839a 

i-Cercopithecus erythrogaster Murie 1866. 

i-Cercopithecus cynosurus 

i-Cercopithecus talapoin 

i-Cercopithecus cephna (C. cephus) 

i-Cercopithecus nictitans J 

i-Cercopithecus callithricus (C. sabaena) Hunter 1861. 

i-Cercopithecus petaurista Ruge 1906. 

-f Cercopithecus sinicus } T/V.UI i oon 

i-Cercopithecus aethiops / 

-\-Cercocebus fuliginosus Bradley 1903. 

i-Erythrocebus patas (Cercopithecus) Ruge 1906. 

-\-Colobus vellerosus Flower 1872. 

i-Theropithecus rueppelli (Gelada) Garrod 1879a. 

-f Papio lestes Mentzer 1929. 

+Papio sphinx Ruge 1906. 



+ Papio maimon. 
-\-Papio anubis . . 



+Papio porcarius. 



Ruge 1906. 
Hunter 1861. 
Ruge 1906. 
Hunter 1861. 
Flower 1872. 
Hunter 1861. 



-\-Papio leucophaeus (Mandrillus) ........................ Sonntag 1922. 

-\-Papio hamadryas .................................... Schrieber 1932. 

-f- Macaco maura ......................................... Murie 1872. 

-\-Macaca brunnea ..................................... Anderson 1872. 

-f Macaco albibarbala (M. silenus) . . . \ -, QC1 

+Macaca mulatta (M. rhesus) .......................... / * 

+M acaca mulatta (M. rhesus) ........................... Linback 1933. 

+M acaca cynomolga ..................................... } 

+ M acaca sinica ........................................ } Ruge 1906. 

+Macaca nemestrina .................................... J 

Pongidae f R uge 1906a. 

+Hylobates lar ....................................... \ Hunter 1861. 

[ Flower 1872. 
+Hylobates lar (H. leuciscus) ........................... Chapman 1900. 

Owen 1830. 
+Pongo pygmaeus ("orang") ......................... \ Sonntag 1924a. 

Ruge 1906a. 



+Pan satyrus ("chimpanzee") .......................... 

+Pan satyrus (Anthropopithecus troglodytes) ............... Sonntag 1923. 

f Flower 1872. 
+Gorilla gorilla ....................................... \ Fox 1930. 

( Ruge 1906a. 
Hominidae 

+Homo sapiens ............................................ A.U-C. 

TABLE 10 
Carnivora 

Canidae 

+Cants familiaris ...................................... Gorham 1936. 

-(-Cants aureus ........................................ Mentzer 1929. 

+Canis lupus ....................................... Macalister 1867. 

+Speothos venaticus (Icticyon) ............................ Flower 1880. 

+Lycaon pictus .............. . / Crisp 1855 

1 Garrod 18<8a. 



1938 GALL BLADDER GORHAM AND IVY 193 

TABLE 10 Continued 
Carnivora (continued) 

Canidae (continued) 

+Otocyon virgatus (Cam's otocyon virgatus) ................. Mentzer 1929. 

+ Urocyon cinereoargenleus (Canis) ........................ Hunter 1861. 

-j-Cuon dukhunensis ..................................... Murie 1872c. 

+Fennecus zerda (Canis) ................................. Hunter 1861. 

+Nyctereutes procyonoides ............................... Garrod 1878a. 

Procyonidae / Crisp 1862. 

+Procyon lotor . ( Macalister 1867. 

+Polos flatus (P. caudivolvulus) .. . { {2^}^ 

+Nasua rufa or narica .................................. Hunter 1861. 

+Bassaricyon gabbi (B. alleni) .......................... Beddard 1900a. 

-f Ailurus fulgens ....................................... Flower 1870. 

+Aeluropoda melanoleuca ..................................... 47432. 

Ursid *' / Hunter 1861. 

+ Unusardos.. \Macalisterl867. 

-1- Ursus maritimus ........................................ Crisp 1862. 

+ Ursus malayanus ........................................ Rex 1888. 

+Melursus ursinus (Ursus labiatus) ....................... Flower 1872. 

Mustelidae 

+Mustfla putorius (Putorius furo) ................... \ Mapa i:, t _ P 18fr 

+Mustela erminea (Putorius) ........................ / Macallst er l b , . 

+Mustela vison ........................................ Gorham 1936. 

+M ellivora sagulata ................................... 1 

+ Taxidea taxus (Meles labradoria) ...................... ( w n OC1 

+Meles meles (M. taxus) .............................. 

+Marles martes (Mustela) ............................. J 

+ Maries pennant i (Mustela canadensis) ................... Martin 1833b. 

+Mydaus marchei (M. meliceps) ........................ \ TT 1 fiP1 

+Grison vittata (Mustela grison vittaius) .................. f a 

+ Tayra barbara (Galera) .................................. Crisp 1862. 

+Helictis subaurantiaca ................................. Garrod 1879c. 

-\-Heliclis personata ................................... Beddard 1905a. 

+Lulra lulra (L. vulgaris) .............................. { cri^fse! 61 " 

+Gulo sp. (Ursus gulo) .................................... Crisp 1862. 

Viverridae 

+ Viverra zibetha ........................................ Hunter 1861 . 

-f Viverra civetta ........................................ Mivart 1882. 

+Suricata suricata (S. telradactyla) ..................... / ' 



+Paguma larvata (Paradoxurus) .......................... Mivart 1882. 

-f Nandinia binotaia .................................... Carlsson 1900. 

Nandinia binotata ..................................... Mivart 1882. 

+Crossarchus obscurus ................................... Martin 1834. 

+Arctictis binturong .................................... Garrod 1873a. 

+Hemigalus derbyanus .................................. Mivart 1882. 

+Galidea elegans ..................................... Beddard 1909a. 



" 1 " 1882 - 

+Cryptoprocta ferox .................................... Beddard 1895. 

Hyaenidae 

+Hyaena brunnea ........................................ Murie 1867. 

+Hyaena vulgaris ....................................... Hunter 1861. 

+Crocuta crocuta (Hyaena) ................... Watson and Young 1879a. 

+Proteles cristata ........................................ Flower 1869. 



194 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 10 Continued 
Carnivora (continued) 

Felidae 

+Felis domestica ........................................ Mivart 1881. 

+ Felis pardcdis ......................................... I /-___ i o fi . ? 

+ Fdis nebulosa (F. macrocelus) ........................... f L 

+Panthtra leo (Felis lea) . . { ggjj/f &. 

+Panthera pardus (Felis leopardus) ....................... Hunter 1861. 

-\-Panthera pardus (Felis pardus) ......................... Mentzer 1929. 

+Panthera onca (Felis) .................................. Martin 1832. 

-\-Lynx caracal (Felis) ................................... Hunter 1861. 

-i-Acinonyx jubala ..................................... Mentzer 1929. 

4-Felis concolor ........................................ Martin 1833a. 

Otariidae 

-\-Eumetopias jubata (Otaria) .............................. Murie 1868. 

-\-Arclocephalus hookeri (Otaria) .......................... Murie 1867a. 

Odobaenidae 

+Odobaenus rosmarus (Trichechus) ........................ Murie 1870a. 

Phocid ^t Hunter 1861. 

+Phoca ntuhna. . 



f Macalister 1867. 
+Phoca groenlandica ............................... { Crisp 1862. 

1 Murie 1870b. 

TABLE 11 
Xenarthra 

Bradypodidae , Hunter 186L 



Bradypus tndactylus. . .......................... s B ur i e t 191 1. 

Bradypus cuculliger (B. cuculli) .......................... Burlet 1911. 

/ Wislocki 1928. 
-Bradypus gnseus .................................. ( Sonntag 1921a. 

Choloepodidae f Hunter 1861. 

+Choloepus didactylus (Bradypus) .................... j a PP 18 o 2 - 

1 Burlet 1911. 
[ Sonntag 1921a. 
+Choloepus hoffmani ................................... Wislocki 1928. 

Myrmecophagidae 

-\-Myrmecophaga jubata ................................... Owen 1854. 

f Burlet 1911. 
-{-Cyclopes didactylus (Cyclothurus) ..................... I Flower 1872. 

( Wislocki 1928. 
+ Tamandua sp ........................................ Wislocki 1928. 

+ Tamandua tetradactyla ................................ Beddard 1909. 

Dasypodidae ( Hunter 1861. 

+Dasypus novemcinctus ................................ ] Crisp 1862. 

( Rapp 1852. 
-\-Dasypus novemcinctus (D. peba) ......................... Owen 1831a. 

-\-Euphractu8 sexcinctus (Dasypus) ....................... | 9-^ n 186^ 

+ Tolypeutes tricinctus ................................... Garrod 1878. 

+ Tolypentes sp ......................................... Murie 1872a. 

+Chlamydophorus truncatus ........................... Macalister 1873. 

Pholidota 
Manidae 

+Manispentadaclyla.. . { Hunter 1861. 

-\-Smutsia temminckii (Manis) ............................ Peters 1852. 

-\-Manis javanica ........................................ Adams 1859. 



1938 GALL BLADDER GORHAM AND IVY 195 

TABLE 11 Continued 
Tubulidentva 

Orycteropidae i j aege r 1837. 
+0rycteropus afer (O. capensis) . . { gonntag 1925. 
+Orycteropus afer (O. afro) F* 1930. 

TABLE 12 
Artiodactyla 

Tayassuidae , Hu nter 1861. 

-Pecan torquatus (Dicotyles) . . . | Macalister 1867. 

Suidae 

+Babirussa alfurus (B. babirus) Flower 1872. 

+Phacochoerus africanus Mentzer 19-29. 

-\-Phacochoerus aethiopicus (P. pallasii) Owen 1851. 

Hippopotamiidae [ Garrod 1879b. 

-{-Hippopotamus amphibius \ Weissberg 1932. 

[ Chapman 1881. 

-Hippopotamus amphibius Chapman 1881. 

Camelidae 

-Corned bactrianus Flower 1872. 

-Lama guanicoe (L. pacos) ... -I c r j sp igg2. 

Lama vicugna (Auchenia sp.) J 

Tragulidae 

+ Tragulus kanchil Hunter 1861. 

+ Tragulus javanicus (T. napu) Macalister 867. 

-\-Dorcatherium aquaticum (Hyomoschus aquaticus) Flower 1867. 

Cervidae 

+Cervus axis Crisp 1862. 

/ Hunter 1861. 
-Cervus axis \ Raven 1936. 

Cervus unicolor unicolor (C. hippelaphus) . \ c r j sp 1862. 

Cervus unicolor moluccensis (C. moluccensis) / 

Cervus unicolor sunnhoii (C. swinhpii) 

Cervus unicolor marianus (C. mariannus) 

Cervus elaphus 



Cervus kukulii . 



Garrod 1877. 



Cervus timoriensis (C. molucensis) 

Cervus duvauceli 

Cervus alfredi 

- Cervus porcinus 

-\-Mazama superciliaris (Cervus) 

-Mazama rufus (Cervus) 

.. , J Crisp . 

-Odocoileus hemionus (Cervus auntus) | Gorham 1936. 

Odocoileus mexicanus (Cervus) \ c r i s p 1862. 

- Alces alces (Cervus) / 

-Alces alces (A. machlis) Watson and Young 18,9. 

Muntiacus muntjac (Cervulus) \ G arro d 1877. 

Af untiacus reevesi (Cervulus reevesi) / 

- Hydropotes inermes Garrod 187 < a. 

-Pudu pudu (Cervus) Garrod 18. 

-Pudu pudu Flower 1875. 

Elaphodus cephalophus J 

Elaphodus cephalophus (Lophotragus michianus) } Garrod 187 /. 

Blastoceros bezoarticus (Cervus campestris) J 

' Garrod 1877. 



+Moschus moschiferus ] Crisp 186 2- 



196 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 12 Continued 
Artiodactyla (continued) 

Giraffidae j Owen 1838b 

1+Giraffa camelopardalis < Garden 1787. 

Owen 1838b. 
Crisp 1862. 
Neuville 1914. 



Giraffa camelopardalis . 



Murie 1872c. 



Chapman 1875a. 

Joly and Lavocet 1845. 

Giraffa camelopardalis (Camelopardalis giraffa) . . < p ox ^929. 
-Giraffa reticulata Lonnberg 1912. 

Antilocapridae , Murie 1870b 

+Antilocapra amencana . . ' Lonnberg 1909. 

Bovidae 
Bovinae 

+Bo8taurus. -\Hunterl861. 

+Bison bison (B. amencanus) J 

+Syncerus coffer (Buffdus c. radcliffei) Mentzer 1929. 

Caprinae 

+Ovis musimon Crisp 1 862. 

-f Copra hircus (C. picta) Garrod 1877. 

* Copra angoriensis (3 specimens) . . Crisp 1862. 

+Hemitragus jemlahicus (Copra jemlaica) Garrod 1877. 

Rupicaprinae 

+ Rupicapra rupicapra Rex 1888. 

-i-Budorcas taxicolor Lander 1919. 

Ovibovinae 

+Oribos moschatus Lonnberg 1900. 

Bubalinae 

Bubalis buselaphus (Antelope) \ c r } s p jggo 

Damaliscus pygargus (Antelope) j 

-4-Damaliscus pygargus (Damalis) Garrod 1877. 

+Connochaetes albojubatus Mentzer 1929. 

+Connochaetes gnu (Catoblcpas) Garrod 1877. 

Cephalophinae , ^ 1QC ., 

/- T i it- / A i i \ i Crisp 18bJ. 

Lephalophus maxima (Antelope) . . . . < Q aT ^ O( i 1377 

Cephalophus monticola (C. pygmaeus) Garrod 1877. 

Cephalophus harveyi keniae Lonnberg 1900. 

Cephalophus melanorrhaeus Lonnberg 1909. 

-Cephalophus natalensis. .-,, \ L5n nberg 1900. 

Cephalophus abyssinicus hinder j 

Cephalophus grimmia (Antelope mergens) Crisp 1862. 

,, , , , .,, . ' Lonnberg 1900. 

-( ephalophus ogilbyi | Lonnberg 1909. 

Oreotraginae 

+Ourebia nigricaudatus (Nanotragus) Garrod 1877. 

+Raphicerus campestris (R. neumanni) < jJp^^r^i 909 

+Neotragus pygmaeus Pocock 1918. 

+Rhyncotragus cavendishi (R. kirkii) Mentzer 1929. 

t Six males, three females, others unsexed. 

Communication with breeders of Angora goats, in this country and abroad, failed to reveal any 
breed in which the gall bladder was absent. The number of specimens indicates that Dr. Crisp prob- 
ably was guilty of mis-statement, rather than dissection of anomalous specimens. 



1938 GALL BLADDER GORHAM AND IVY 197 

TABLE 12 Continued 
Artiodactyla (continued) 
Bovidae (continued) 

Reduncinae /-,;, i9 

+Redunca arundinum (Antelope tsabelhana) ............... ... crisp lb. 

+Kobus defassa ....................................... Mentzer 1929. 

4-Kobus ellypsiprymnus ................................... Smith 1! 

S TsaT0a tartarica ........................................ Murie 1870c. 

Antelopinae ,, . , Q< , 9 

+Antelope cervicapra (A. bezoartica) ...................... v" 8 ^ *$?* 

J Crisp lobJ. 

-\-Gazella dorcas (Antelope) ............................. ^ Garrod 1877. 



+GaUa bennetti (Antelope) .. ... -CrP 1862. 



+Gazella granh ...................................... \ Garrod 1877. 

+Gazella subgutturosa .................................. 1 

+Gazella muscatensis ..... . . . . 1 G arro( j 1877. 

-f Gazella rufifrons ..................................... 

+Gazella arabica ...................................... J 

+Antidorca marsupialis (Antelope euchore) ............... ... .crisp I8t>-. 

+Litfiocranius walleri ................................. Lonnberg 1900. 

Oryginae r-, 

+Oryx leucoryx (Antelope) .............................. V 

J Crisp 
+Addax naso-maculatus ............................... \ Garrod 1877. 

Tragelaphinae r crisp 1862. 

+ Tragelaphus scriptus (Antelope) ....................... | Garrod 1877. 

+ Strepsicerus strepsicerus (S. Kudu) ...................... Garrod 1877. 

+ Taurotragus oryx ..................................... Mentzer 1929. 

+ Taurotragus oryx (Antelope oreas) ....................... cusp iw. 

+ Taurotragus oryx (Oreas caana) .......................... Crarroa 18 / /. 

+Boselaphus tragocamelus (Antelope picta) ................. ^ unte j j^i' 

+Boselaphus tragocamelus (Portax picta) .................. XT "n 1007' 

+Limnotragus gratus (Tragelaphus gratus) ................. Neuville ly/. 

+ Tetracerus quadricornis (T. subquadricornatus) ............. Garrod 18 / /. 

TABLE 13 
Sirenia 

+ Trichecus manatus (Manatee americanus) ................ - Murie 1870. 

+ Trichecus manatus (Manatus inunguis) ................. Beddard 189 1. 

-+Trichecus manatus ("Manatee") ...................... Chapman 1875. 

Dugongidae 

+Dugong australis (Halicore sp.) .......................... ^> wen 1] 

Stellerl749. 



Hyracoidea 
Procariidae 

+Procaria capensis 



Mentzer* 1929 
Raven 1936. 
Crisp 1862. 

-Procavia capensis i f}^ er * .. Q . 

I Huxley 1872. 

t Martin 1835. 

Mentzer examined two specimens, one with a gall bladder fined with liver flukes. He notes 
that the gall bladder, with a central and two lateral sacs, as reported by Owen and Macalister, ma_y 
be found only as a dilation or a lateral pouch of the common duct, at the point where the four hepatic 
ducts join it. 

** Flower says that this animal has no gall bladder, but in some specimens the common duct 
dilated to great size. 



198 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 13 Continued 
Proboscidea 

Elephantidae f Hunter 1861. 

Elephas maximus (E. indicus) \ Crisp 1862. 

1 Forbes 1879. 

Loxodonta africana (Elephas africanus) Forbes 1879. 

/ Eales 1929. 

- Elephant <[ Chapman 1875b . 

TABLE 14 
Perissodactyla 
Tapiridae 

Tapinis terrestris (T. americanus) Crisp 1862. 

- Tapirus indicus Parker 1882. 

- Tapiru* indicus Murie 1871b. 

Rhinocerotidae 

D , . . . f Beddard 1887. 

- Rhinoceros umcorms < Q wen 1350 

Rhinoceros sumatrensis (Ceratorhinus) Garrod 1873. 

Equidae 

Equus caballus 1 

Equus asinus [ Hunter 1861. 

Equus zebra j 

-Equus burchelli 44391. 

Equus quagga Mentzer 1929. 

TABLE 15 
Cetacea 
Mysticeti 
Balaenidae 

Balaena mysticetus Hunter 1840. 

Balaenopteridae , *. -\oa-i 

- Balaenoptera acutirosfrata (B. rostrata) { Eschricht 1849 

-Megaptera longimana (M. boops) { gajg.^ ^ 4 a 9 Calister 1868! 

Odontoceti 
Platanistidae 

Platanisla gangetica Anderson 1879. 

Physeteridae 

Physeter catodon (P. macrocephalus) Hunter 1840. 

Ziphiidae 

Ziphiorrhynchus cryptodon(l] Burmeister 1866. 

Hyperoodon ampullatus (Delphinus bidens) Hunter 1861. 

Delphinapteridae 

Delphinaplerus leucas (Belugia catodon). . . .Hepburn and Waterson 1901. 



- Monodon monoceros . . { Hjjjjj }^0. 

Delphinidae 

Neophocaena phocaenoides (Neomeris) Chi Ping 1926. 

Phocaena phocaena (Delphinus) Hunter 1840. 

-Phocaena phocaena (P. communis) { ggJS^Ji Wate r S on 1901. 

Orca gladiator (Delphinus orca) Turner 1899. 

Orcella brevirostris (Orcella) Anderson 1879. 

( Gulliver 1853. 

Globiocephalus melaena (G. melon) <^ M 91 "! R^ 

[ Weissberg 1932a. 



1938 GALL BLADDER GORHAM AND IVY 199 

TABLE 15-C<mhued 



Cetacea (continued) 
Odontoceti (continued) 
Delphinidae (continued) 

:i$r&f' "*";. : : } ***** > 

Tursiops truncatus (Delphinus tursio) .................... Hunter 1861. 



+Gramphidelphis risii (Globiocephalus risii) ............... Williams 1838. 

Gramphidelphis risii (Grampus rissoanus) ................ Murie 1871b. 

Rodentia TABLE 16 

Aplodontiidae 

- Aplodonlia rufa .................................. Hall, MVZ 22623. 

Sciuridae ( Crisp 1862. 

+Sciurus vulgaris ..................................... { Rex 1888. 

I Cuvier 1835. 

Sciurus carolinensis ..................................... Crisp 1862. 

+Sciurus carolinensis (S. cinereus) ........................ Hunter 1861. 

Sciurus indicus (S. maximus) r 



+ Tamias striatus ........................................... < 1544 

-\-Marmota marmota (Arctomys) .......................... .Hunter 1861. 

+Marmota marmota (Arctomys alpinus) ............... | MTcalister'lSe?. 

+ Marmota monax ("Marmottes de Canada") ............... Cuvier 1835. 

+Citellus tridecemlineatus ................................ Higgins 1928. 

+Citellus suslicka ("Spermophile souslick") ................ Cuvier 1835. 

\-Glaucomys volans (Pteromys volucella) ................... Yarrell 1831a. 

+Glaucomys volans (Sciuropterus volucella) ................. Hunter 1861. 

+Glaucomys volans (Sciurux volucella) ....................... Crisp 1862. 

+Glaucomys volans ("Hassapan") ....................... } r<uv\er 1835 

Petaurista petaurista ("Le grand ecruil volant de Java"). / 

Castoridae , Macalister 1867. 

+Castor fiber .. -\Crispl862. 

-\-Castor canadensis ..................................... Hunter 1861. 

Heteromyidae 

-Perognathus fallax ........................................... 16190. 

Geomyidae 

- Thomomys bottae ...... . \ Gorham i 936 . 

Geomys our sari us ................................... ) 

Anomaluridae 

+Anomalurus pelii. . - 1 AUton 1875- 

-\-Anomalurus frasert ................................... ) 

Pedetidae 

-Pedetes caffer ............................. - Parsons 1898. 

Pedetes caffer (Helamys capensis) ........................ Hunter 1861. 

-Pedetes caffer (Forster's jerboa) .......................... Cuvier 1835. 

Myoxidae 

. / Cuvier 1835. 

+Muscardtnus avellenanus. . . j Macalister 1867. 

+G(t glix ("loir") . . . . ..... . 1 Cuvier 1835- 

+Glis nitedula (?) ("lerot") ........................ 

Dipodidae ( Hunter 1861. 

+Dipus saggita ...................... { Macalister 1867. 

I Duvernoy and Lereboullet 1840. 



200 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

TABLE 16 Continued 
Rodentia (con/inued) 
Dipodidae (continued) 

+Dipus gerboa (D. mauritanicus) Duvernoy and Lereboullet 1840. 

+ Jaculus loftusi 42459. 

-\-Zapus hudsonius 18804. 

+Zapus hudsonius (Dipus americanus) Cuvier 1835. 

Spalacidae 

-\-Myotalpa aspalax (Siphneus myospalax) Milne-Edwards 1874. 

\-Spalax sp Macalister 1867. 

+ Spalax typhlus (M us) Cuvier 1835. 

+Rlnzomy8 badius .} Anderson 1879 . 
+Rhizomys pruinosus / 

Muridae 
Cricetinae 

Cricelus cricetus Macalister 1867. 

Cricetus cricetus ("Hamster") 

Cricelulus migratorius (Mus alticus) . 



Cricetulus migratorius (Mus accedttla) 



Cuvier 1835. 



Cricetulus migratorius (Mus phaeus) 

Cricetiscus sungarus (Mu$). 

*Reithrodontomys longicauda ................................... 16173. 

-\-Peromyscus leucopus ......................................... 15484. 

Lophiomyinae 

Lophiomys sp ......................................... Flower 1872. 

Mi f r f inae , / Macalister 1867. 

+Armcola amphibia. . { Hunter 18gl 

+Dicrostonyx richardsonii ("Lemming de la baie 

d'Hudson") ...................................... I /-...__ 1 00= 

+Ellobins talpinus (Mus) .............................. 6 ^ 

+Ondatra zibethicus ("Ondatra") ....................... J 

+Ondatra zibethicus ("Desman") ......................... Hunter 1861. 

-\-MicrotuK ochrogaftter ......................................... 15486. 

Murinae 

Rattus norvegicus (Mus decumanus) ...................... Hunter 1861. 

+Pelomys fallax ......................................... Peters 1852. 

+Mus musculus 



+ Leggada bellus (Mus minimus) ........................... Peters 1852. 

Apodemus ayrarius (Mus) ............................ \ /-.._ _ -, 8q c 

- Micromys minutus (Mus) ............................. / ^ 

+Steatomys pratensis (S. edulis) ........................... Peters 1852. 

+Arvicanthis abyssinicus ....................................... 21302. 

+Arvicanthis barbarus (Mus) ............ Duvernoy and Lereboullet 1840. 

Otomys nyikae .............................................. 21274. 

+Otomys brantsii (Euryotis) ............................... Smith 1849. 

Gerbillinae 

+Meriones shawi ("Gerbille de Shaw") .................... Cuvier 1835. 

+Meriones shawi (Gerbillus shauni) ...... Duvernoy and Lereboullet 1840. 

+Gerbillus pygargus ("Gerbille du Senegal") ................ Cuvier 1835. 

+ Tatera leucogasler (Meriones) ............................ Peters 1852. 

Hydromyinae 

Hydromys chrysogaster ................................. Windle 1887. 

Bathyergidae 

+Myoscalops argenteo-cinereus ............................. Peters 1852. 

+ Rathyergus sp ....................................... 1 

+Georychiis capensis (Orycterus) ........................ / " bl - 

* In one n( four individuals the organ was present. 



1938 GALL BLADDER GORHAM AND IVY 

TABLE 16 Continued 
Rodentia (continued) 
Hystricidae 

... .r* i*. 



+Hystrix cristata ("Pore-epic") ........................... Cuvier 1835. 

-Hyslrix africafaustralis .............................. ; Pete l^p- 

. , / Parsons 1894. 

+Acanthion javamcum (Hystnx jamnica) . " ' \ Mivart 1882a. 

Erethizontidae 

-Ereihizon dorsalis ................................... Mivart 1882a. 

Macalister 1867. 



Coendou prehensilis (Hystrix) 
Dasyproctidae 



4-Dasyprocta aguti 



Cuvier 1835. 

Macalister 1867. 

Crisp 1862. 

Jones 1834. 

+Dasyprocta antillensis or albida (D. cristata) Mivart and Murie 1866. 

i Owen 1831. 
+Myoprocta acouchy (Dasyprocta) ^ Hunter 1861. 

Caviidae f Hunter 1861. 

+Cavia porcellus \ Macalister 1867. 

i Higgins 192(. 
f Stark 1934. 
j Macalister 1867. 
-\-Hydrochoerus hydrochaens (H. capybara) < Hunter 1861. 

( Crisp 1862. 

+Cuniculus paca (Coelogenys) i?^?-* 6 

+Cuniculus subniger (Coelogenys) M artin 

+Dolichotis patagonica Beddard 1 81. 

Chinchillidae 

+Chinchilla laniger -I Bennet 1833. 

+Lcgidium viscaccia (Lagotis cuneri) j 

+Lagidium riscaccia (Lagostomus trichodactylus) Owen 

Capromyidae 

+Capromys pilorides (Isodon) 

., -j //- f -\ <-- 

+Capromys pilondes (C. fourmen) .... . < Cuvier 1835. 

/ Kuhl 1820. 
+Capromys melanurus - | Dobson 1884. 

Octodontidae 

+Octodon degus (O. cumingii) Martin I<w6b. 

- Echimys sp Cuvier 1 3o. 

+Pectinator spekii era ai \' 

+Ctenomys torquatus 

-\-Myocastor coypus (Myopotomus) Martin 

Ctenodactylidae 

-\-Ctenodactylus gundi (C. massonii) Yam Ji. 

Petromyidae Q 

-Petromys typicus 4y - 

Thryonomyidae IQ.TQK 

-\-Thryonomys swinderianus (Aulacodus) .Lrarroa IMD. 

Leporidae ( Crisp 1862. 

+Lepus timidus \ Hunter 1861. 

+Lepus californicus Gorham 1936. 

-\-Oryctolagus cuniculus (Lepus) .. Hunter 1 ( - ; . 

Ochotonidae 

-f Ochotona pnnceps .MVZ 916. 

* Smith figure* the liver. No gll bladder is shown. He offers no discussion. 



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204 FIELD MUSEUM OF NATURAL HISTORY- ZOOLOGY, VOL. XXII 

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1938 GALL BLADDER GORHAM AND IVY 205 

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206 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

GARDON (GORDON?) 

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1876. On a Peculiarity in the Carotid Artery, and Other Points in the Anatomy 
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1879. Notes on the Visceral Anatomy of the Tupaia of Burma. Proc. Zool. 
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GARROD, A. H. and DARWIN, F. 

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1938 GALL BLADDER GORHAM AND IVY 207 

GUNTHER, A. 

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1938 GALL BLADDER GORHAM AND IVY 209 

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210 FIELD MUSEUM OF NATURAL HISTORY ZOOLOGY, VOL. XXII 

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